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A REVISION OF THE BRACHIOPOD SUBFAMILY
KINGENINAE ELLIOTT

By E. F. OWEN

CONTENTS

I. Introduction ........

II. Acknowledgments .......

III. Techniques ........

IV. Previous research .......

V. Stratigraphical range and geographical distribution .

VI. Morphology and structure ......

VII. Shell structure ........

VIII. Systematic descriptions ......

IX. Conclusions .........

X. References .........

Page

SYNOPSIS

This paper gives an account of previous research on the Dallinidae and comparison is made
between the early loop development stages of both fossil and living representative genera.

The stratigraphical range and geographical distribution is given in fair detail with correlation
tables for European, Asian and American Cretaceous horizons.

The family name Kingenidae nom. transl. for Kingeninae, Elliott 1948 is suggested and
includes species of Kingena, Zittelina and Belothyris, It proposes NEOTYPES for K, arenosa,
K. mesembrina, K, lima and K. pentagulata and Belothyris pseudojurensis. LECTOTYPES
selected for Kingena lemaniensis and Zittelina orhis are redescribed in the light of present
knowledge.

A new genus, Waconella, is erected for the North American species wacoensis, formerly assigned
to the genus Kingena and now shown to be closely related to the Tertiary and Recent genus
Laqueus.

I. INTRODUCTION

The purpose of the present research has been to investigate the early developmental
stages of the brachial loop and cardinalia of certain Terebratellacean brachiopods
within the Cretaceous and to show how these characters can confirm or justify the
position of the genera in the classification and, at the same time, to compare and
contrast the fundamental characters of mature forms and to illustrate their possible
relationship to similar structures in Jurassic progenitors and Recent descendants.

The suborder Terebratellidina is divided into three superfamilies, the Crypto-
nellacea, Zeilleriacea and the Terebratellacea. One of the most important families
within the Terebratellacea is the Dallinidae, as set out in the Treatise of Invertebrate
Paleontology. The present work revises some of the genera within the family.

THE BRACHIOPOD SUBFAMILY

particularly those previously assigned to the subfamily Kingeninae. As a result of
the present study, the kingenids have been raised to family level, parallel to the
Dallinidae.

Apart from the well known works of Deslongchamps and Davidson in the last
century and later, Muir-Wood, very little research work has been done on the Mesozoic
Terebratellidina as a whole and hardly anything is known about the fossil Dallinidae
in particular. Recent interest in the possible origin of this family, however, has
been shown by the Russian workers Babanova and Dagis, who have illustrated early
dallinoid type loop attachments in species of Aulacothyris from the Jurassic of the
Crimea. Valuable contributions to our knowledge of the Terebratellidina have been
made by the searching enquiries of these two authors but in both cases reference is
made to western European species of Aulacothyris while their comparative material
has been drawn solely from eastern European sources.

Many gaps in our knowledge of the early loop development, the vertical range and
the geographical distribution of the Terebratellidina will have to be filled before any
real support can be given to such drastic steps as suggested by Dagis in the taxo-
nomic position of some of the genera he examined.

Material for this investigation has been difficult to obtain, particularly representa-
tive species of the genexdiTulipina, Belothyris and Zittelina. It is for this reason that
a more exhaustive study of species of the comparatively common genus Kingena has
been made.

A complete monograph of the Kingeninae has not been possible as, in some cases,
no comparative material has been available for further study and the original
illustrations of some species has not always justified their reference to this subfamily.

II. ACKNOWLEDGMENTS

I wish to express my appreciation to the numerous people who have so kindly given
me assistance during the course of this work.

I am particularly indebted to my colleagues Drs. L. R. M. Cocks and C. H. C.
Brunton for their helpful suggestions and amiable company ; to the Keeper of
Palaeontology British Museum (Natural History), for permission to work on the
collections in his charge ; to Drs. N. J. Morris, R. P. S. Jefferies and G. F. Elliott, of the
same department, for their wise counsel ; to Dr. W. J. Kennedy, Oxford University ;
to Dr. Colin Forbes, Sedgwick Museum, Cambridge ; Mr. C. J. Wood, Institute of
Geological Sciences ; Dr. P. J. Coleman, Department of Geology, University of
Western Australia ; to Dr. Margaret Pinder for helpful discussion on Cretaceous
brachiopod faunas ; to Dr. E. Lanterno, Musee d'Histoire Naturelle, Geneva ; to
Mile. Anny Dhondt, Inst. roy. Sci., Belgique, Brussels ; to Dr. W. T. Dean, Geological
Survey of Canada ; to Messrs. Peter Green and Tim Parmenter, British Museum
(Nat. Hist.), for most of the photographs ; to Dr. R. Goldring, University of Reading,
for helpful advice and to Mr. Brian Martin and members of the staff of the Electron
Microscopic Unit, British Museum (Nat. Hist.).

My very sincere thanks are due to Professor P. Allen, University of Reading, who
supervised this work and who generously helped in many other ways ; to Professor
Alwyn Williams for friendly discussion in the early stages of the work ; to Dr. A. J.

KINGENINAE

Rowell of the University of Kansas, and to my wife Mary, for her continued en-
couragement and supreme tolerance.

There are many others who have not been mentioned above and to whom I also
owe my thanks.

III. TECHNIQUES

The classification of the brachiopoda depends very largely upon an intimate know-
ledge of the finer structures of the cardinalia and the brachial loop. In fossil
genera this knowledge can be obtained in several ways. Dissection of the brachial
apparatus from the surrounding matrix infilling the shell is sometimes possible pro-
viding that the matrix is sufficiently soft or is not recrystalline. Various methods
have been described which make this procedure less dependent upon a high degree of
technical skill. One of these is the use of an ultrasonic waterbath which removes
soft matrices from brachial loops with the minimum of damage to the structures.
This method has been freely used here for specimens from the Upper Chalk.

Where dissection is not possible, transverse serial sections are often used. This
method described by Muir-Wood (1934), with minor modifications, has been em-
ployed for most of the specimens from either limestone or hard clay localities.

For the sake of fair comparison, this method has been used for more critical study
of cardinalia and hinge structures in relation to brachidia. To aid differentiation of
these characters in the case of specimens from the Upper Chalk, a technique involving
the use of sugar solution described by Owen (1955) has been employed.

Where comparison has been made with Recent species, examples of dried specimens
have been filled with an artificial matrix composed of plaster of Paris and asbestos
powder and serial sectioned in the same way as the fossil forms.

Use has also been made of the Cambridge 'Stereoscan’ electronmicroscope for the
investigation of shell mosaic of both Recent and fossil species and also for the pus-
tulate shell ornament.

IV. PREVIOUS RESEARCH

Since the publication of the Treatise on Invertebrate Paleontology (1965), the
DaUinidae have become a well established family within the suborder Terebratel-
lidina. The essential differences separating this family from other families within
the suborder can best be illustrated by comparison of the very early growth stages in
the development of the brachial loop.

In the Recent Terebratellidina, some of the more complex stages of metamorphosis
in the ontogeny of the loop have already been described by Friele (1876, 1877),
Deslongchamps (1884) and Fischer & Oehlert (1892). Basing his classification on
major differences in these early stages, Beecher (1895), split off three subfamilies
from the main terebratelloid group. These were the Dallininae, Magellaniinae and
Megathyrinae and were later discussed and revised by Thomson (1937) who added the
Platidiinae, Muhlfeldtiniae and Laqueinae. In 1940, Allan proposed the super-
family Terebratellacea with five new families, among which, the Dallinidae, had two
subfamilies namely, the Dallininae and Laqueinae.

The new set of terms for the dallinoid growth stages which arose from the work of

THE BRACHIOPOD SUBFAMILY

the earlier authors was also discussed by Thomson (1927) who revised some of the
nomenclature. Fischer & Oehlert (1892) had introduced the term centronelliform
for the earliest loop development stage in the Dallinids ; Beecher (1895) replaced this
term by platidiform in his classification. This was once again replaced by Thomson
(1927) who used the term pre-ismeniform for this early stage. A further correction
was made by Elliott (1947) who pointed out that the loop in Ismenia pectunculoides
(Schlotheim), the type species of Ismenia from the Upper Jurassic of Germany, does
not correspond with what is known as ismeniform stage of loop development of the
Recent Dallinids and proposed the alternative names of pre-campagiform and cam-
pagiform from the Recent genus Campages to replace the terms pre-ismeniform and
is meniform as used by Thomson (1927).

Most of the terminology and classification surrounding the dallinoid brachiopods,
therefore, has been developed from the study of Recent genera. Comparatively
little is known about the Mesozoic Dallinidae and little advance has been made on the
work of earlier authors such as Deslongchamps and Fischer & Oehlert.

Over the past twenty years, however, further contributions to the classification
have been made by Elliott (1947) who described the genus Gemmarcula from the
Lower Cretaceous, Upper Aptian of Faringdon, Berkshire, illustrating his description
with a series of carefully prepared dissections of early dallinoid brachial loop stages.
For this genus Elliott proposed the subfamily Gemmarculinae.

Later Elliott (1948) described and illustrated a series of early loop stages in the
genus Hamptonina from the Great Oolite of Hampton Down, Bath. These show the
pre-campagiform and campagiform stages to perfection and the specimens are again
used in the present work, PI. ii, figs. 1-3, for comparison with early growth stages in
the brachial loops of specimens of Kingena from the Upper Chalk.

In 1948, in a paper discussing the evolutionary significance of the brachial develop-
ment of terebratelloid brachiopoda, Elliott proposed the subfamily name of Kinge-
ninae for the Cretaceous genus Kingena Davidson, 1852. To this has been added
Zittelina, erected by Rollier (1919) for Terehratula orhis Quenstedt (1871) from the
Upper Jurassic of Wurttemburg, Germany ; Belothyris, described by Smirnova
(i960) from the Lower Cretaceous of the southern Caucasus and Tulipina which
Smirnova (1962) described from the Lower Aptian of Georgia.

The proposal of a new subfamily, Belothyrinae for the genus Belothyris was made
by Smirnova (i960) but this was not adopted by Elliott in preparation of the section on
Dallinidae for the Treatise on Invertebrate Paleontology (1965).

Further details concerning the relationship of the above genera and a comparison
of internal structures in both Recent and fossil representative species of the Dallinidae
is given in the foregoing sections on Morphology and Structure and in the Systematic
Descriptions of the present work.

V. STRATIGRAPHICAL RANGE AND GEOGRAPHICAL DISTRIBUTION

The accepted stratigraphical range for the subfamily Kingeninae is that quoted in
the '' Treatise ” (1965) as Upper Jurassic to Cretaceous. Of the four genera so far
included in the subfamily, the genus Zittelina of Rollier (1919), occurs in the Upper
Jurassic, Weiss Jura of Wurttemburg, Germany from whence the type species.

KINGENINAE

Terebratula orbis Quenstedt (1858), was originally described. This genus also occurs
in the hard limestone facies of the Tithonian of Stramberg, the Stramberger-
Schichten '' of Czechoslovakia and from a similar horizon at Card and Herault,
France. In 1905, Krumbeck extended the distribution of the genus to the eastern
Mediterranean, describing two new species, Z. latifrons and Z, triangularis, and
recognizing three of QuenstedFs species, Z. orbis, Z, gutta and Z. cubica from the
Upper Jurassic of the Lebanon. He mistakenly referred these to the Cretaceous
genus Kingena, but they are here referred to the genus Zittelina.

The genus Belothyris was described by Smirnova (i960) from the Lower Cretaceous
of northwest Caucasus. Examples of topotype material of three species, B. plana,
the type species, B. regularis and B. marianoviensis have been kindly presented
to the British Museum (Natural History) by Dr. Smirnova and are figured here PI. 10,
figs. 3, 4 & 5. These specimens have been used for comparison with specimens
collected from the Lower Cretaceous of Sheik Budin, southeast of Pezu, N. W.
Frontier Province, Pakistan. The comparison is favourable and establishes,
without doubt, that the two forms are congeneric, but more material from both
Russian and Pakistani localities is needed before any realistic comparison of species
can be made.

In her original description of the genus Belothyris, Smirnova suggested that '' Zeil
leria ** pseudojurensis described by Leymerie (1842 : 12) from the Hauterivian of
MaroUes (Aube), France, might belong to her genus. As a result of her suggestion,
an investigation of Leymerie's species pseudojurensis has been carried out during the
course of the present research with the result that this species can now be referred to
the genus Belothyris Smirnova.

Belothyris pseudojurensis (Le3mierie) is widely quoted from Neocomian beds in
France and Switzerland, occurring in the radiatus Zone of the Lower Hauterivian at
Le Landeron, and Neuchatel, Switzerland and from the type locality, MaroUes and
from Auxerres (Yonne), Vendeuvre and Avalleur, France.

No duplicate specimens of Tulipina which Smirnova (1962) described from the
Lower Aptian of Georgia have been obtained for the present study but passing
reference is made to the internal characters of the type species, Terebratula koutaisen-
sis de Loriol from the Hauterivian of Switzerland and is discussed in the section on
Morphology.

Although the geographical distribution of the Kingeninae can be quoted as world-
wide, the records show this to be due in the main to citations of the Cretaceous genus
Kingena. Not all the records appear to be authentic, but where possible, duplicate
material has been obtained and studied and is either referred to or figured here.

European records of Kingena are chiefly from British, Belgian, French and German
localities. In France, the type species Terebratula lima was originally described by
Defrance (1828) from an undoubted Upper Chalk locality in the Beauvais district of
Normandy. Unfortunately, no specimen identified as T. lima by that author has
survived and the vague reference to craie de Beauvais '' has led to somewhat wild
speculation as to its stratigraphical position. In addition to Defrance's description,
d'Orbigny (1847 : 108) described a specimen from the Upper Senonian of Chavot,
(Marne), as Terebratula hebertiana. This species is regarded here as a synonym of

THE BRACHIOPOD SUBFAMILY

Fig. I. Sketch-map showing global distribution of the Kingenidae.

KINGENINAE

Woodward's Kingena pentangulata described from the Upper Senonian, Belemnitella
mucronata Zone of Norfolk in 1833.

d’Orbigny made further reference to a species of Kingena in the same publication
(1847 : 98, pi. 512. fig. 1-5) with his description of Terebr alula lima which he quoted
from the Cenomanian of Le Havre. This reference is discussed under the description
of Kingena arenosa (d' Archaic) p. 55, to which species d'Orbigny's specimen ha?
been assigned.

In Belgium, species of Kingena have been quoted from Cenomanian localities in
the districts of Tournai and Montignies-sur-Roc, d' Archaic having described Kingena
from the Tourtia of Gussignies.

From the Upper Chalk of the Mons and Ciply districts, two species are compara-
tively common fossils. Kingena hlackmorei sp. nov. occurs in the Craie de Trivieres
at approximately the same horizon as the Gonioteuthis quadrata Zone in Wiltshire
from which it was originally collected and is probably present in the base of the
Craie d’Obourg. It certainly occurs in the conglomerates which separate the two
beds and are known as Conglomerats d^Obourg.

By far the commonest species of Kingena occurring in the Belgian Upper Chalk is
the British species Kingena pentangulata (Woodward) which occurs at Harmignies
in the Craie de Nouvelles and at Obourg in the Craie de Spiennes and Craie d'Obourg
(see Table i p. 36).

German records, on the other hand, are not so plentiful, being those of Geinitz
(1850) and Schloenbach (1866, 1867, 1868) for species from the Middle and Upper
Chalk of Westphalia.

Roemer (1840) described and figured many brachiopods from various horizons
within the Cretaceous of north Germany. Among these, Kingena pectoralis from the
Hilsconglomerat near Essen, was described (p. 42) and figured (pi. 7, fig. 19 a, b, c.)
as Terebr alula pectoralis. The term Hilsconglomerat, as used by Roemer (1840) can
be interpreted to cover beds ranging in age from Lower Noecomian to Cenomanian
(see Stolley, 1908). Roemer's specimen could have come from any one of those
horizons. Furthermore, the specimen figured by Roemer on pi. 7 is shown as an
elongate-pentagonal specimen which could very well have orignated from the Albian
or Cenomanian. The species Kingena pectoralis (Roemer) is, therefore, regarded here
as nomen dubium.

Outside Europe, records of Kingena extend to Greenland, Canada, north and south
America, Madagascar, Africa, southern India, Tibet and Australia.

Of these records, not all are reliable. The species described by Frebold (1935) as
Kingena groenlandica from the Aptian of East Greenland is not referable to this
genus. As figured and described by Frebold, it shows none of the diagnostic charac-
ters of the genus and may possibly belong to Tamar ella Owen 1965.

The species described by Whiteaves (1903) as Kingena occidentalis from the Upper
Chalk of Canada, bears only a superficial resemblance to the genus and must be left
in doubt until more material from this locality is available. The short septum and
lack of any obvious dental plates suggests a possible relationship with a terebratellid
recently described as Kingenella from the Maastrichtian of Poland by Ewa Popiel-
Barczyk (1968).

Table i

THE BRACHIOPOD SUBFAMILY

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NVINON3S

Showing correlation of British and western Belgian Upper Chalk beds. (Taken partly from Peake & Hancock 1961).

KINGENINAE

References to Kingena wacoensis (Roemer), which was originally described by F,
Roemer (1852 : 81) from the Lower Cretaceous of Texas, U.S.A., have been numerous
and quotations of this species frequently appear in faunal lists of Lower Cretaceous
fossils from Mid West and southwestern localities of Central U.S.A. and from
Mexico.

Although this species is already well entrenched in American literature under the
generic title of Kingena, recent investigation of this form in preparation for the
present work has revealed that it belongs to a completely different generic stock from
Kingena and is described here as Waconella gen. nov. Lack of plentiful material from
all the known localities and horizons has prevented a thorough investigation of the
genus but it is hoped in future to follow up this study with further investigations of a
more concentrated nature.

The occurrence of Waconella wacoensis (Roemer) in the Comanchean Series is dis-
cussed under the description of the genus but a correlation table, Table 3 p. 40, is
included here to give some guide as to the relative age of the beds in terms of European
stratigraphy.

Richter (1925) referred a brachiopod from the Upper Cretaceous of Tierra del
Fuego, Chile, South America, to Kingena lima but the specimens he figured (pi. 9,
figs. 31, 32) show the internal moulds of a small cuboidal brachiopod species with a
short median septum and carinae bordering a shallow sulcus on the brachial valve.
It is a fragmentary specimen with a small incurved umbo and minute foramen and is
almost certainly not referable to the genus Kingena.

African records are those of Ficheur (1900) who recorded Terehratula lemaniensis
Pictet & Roux, here referred to Kingena, from the Lower Albian of Algeria and
Drot (1953) who referred to a Kingena species from the Cenomanian of Tunisia.

In addition, Muller in Bornhardt (1900) described Kingena schweinfurthi from the
Lower Cretaceous of Ntandi, Tanzania but, to judge from the specimen which he
figured (pi. 25, fig. ii), the species would appear to belong to some genus other than
Kingena. The beak characters show a small foramen in a slightly produced sub-
erect umbo with no obvious sign of beak-ridges. No reference is made in the descrip-
tion (p. 548) to external ornament and the figured specimen is seemingly without a
clearly marked median septum in the brachial valve. Although the whereabouts of
Muller's specimen are not known, the species occurs in beds containing “ Trigonia "
swartzi Muller (see Aitken 1956) which has a limited range of Valanginian to Lower
Aptian age. It is conceivable that Muller’s specimen may be referable to the genus
Belothyris from the Hauterivian.

Collignon (1950 : 28) referred a specimen collected from the Albian of Madagascar
to Kingena asperulina Stoliczka which was originally described (Stoliczka, 1872)
from the Utatur beds of Southern India. The specimen which Collignon figured
(pi. 3, figs. 15 a, b.) certainly bears more than a superficial resemblance to the
specimen figured by Stoliczka (pi. 7, fig. 14) and these forms may well be conspecific.
The age of the Utatur beds is given as Upper Albian to Cenomanian in the Lexique
Stratigraphique International (see Table 2 p. 38).

Sahni (1939 : ii) considered that the specimen which he described and figured
(pi. I, figs. 4, 5) from the Pezu district, Pakistan, should be referred to Kingena,

THE BRACHIOPOD SUBFAMILY

3 «

although he admitted that it showed no evidence of pustulate ornament of the shell.
It is quite hkely that K. punjabica belongs to another genus, possibly Belothyris, but
the information regarding its exact stratigraphical origin is vague.

The reference by Muir-Wood (1930 : 34) to two species, K. spinulosa (Davidson &
Morris) and K, tumida Muir-Wood to the Gault of the Samana Range increases the
geographical range of Albian records of the genus to India.

Spengler (1923, pi. i, fig. 13 a, b, & 14) described and figured a crushed specimen
under the name of Waldheimia species from the Upper Chalk of Tharia Ghat, Assam.
The specimen figured by Spengler bears a distinct resemblance to Kingena black-
morei sp. nov. from the Gonioteuthis Zone of Wiltshire.

Table 2

12 ;

" DANIAN
MAASTRICHTIAN
CAMPANIAN

SANTONIAN

CONIANCIAN

TURONIAN

CENOMANIAN
Ur. ALBIAN

I ARRYALUR
J GROUP

I TRICHINOPOLY
J GROUP

\ UTATUR
/ GROUP

Rough correlation of Cretaceous Beds of Southern India with European stages

It is probable that the specimen described by Douville (1916) from the Maastrich-
tian of Tibet as Kingena hebertiana d'Orbigny is an undescribed or new form. It
may, however, be referable to K. shalanurensis Stoliczka from the Arryalur beds of
Southern India.

Further records of Kingena from India were made by Stoliczka (1872) who, apart
from K. granulifera mentioned above, described two further species namely, K.
asperulina from the Utatur group and K. shalanurensis from the ? Trichinopoly
group. These forms are described and discussed briefly in the systematic section
here.

Etheridge's citation of Magas mesembrinus from the Upper Cretaceous of Western
Australia was discussed and revised by Elliott (1952) in a description of some of the
brachiopods from the Gingin Chalk who referred the species to the genus Kingena.

The age of the Gingin Chalk has been the subject of much discussion over the past
fifty years and Glauert (1910, 1926) gave a good general account of its position at
that time. Withers (1924, 1926), following the discovery of the crinoids Uinfacrinus
and Marsupites, pointed out that these fossils occur as horizon markers in the Upper
Cretaceous of Western Europe. More recently McWhae, Playford, Lindner, Glen-
ister and Balme (1958) and Belford (1958) have discussed the position of the Gingin
Chalk and the Toolonga calcilutite in relation to microfaunas, particularly of ostra-
coda.

An unpublished record of a Kingenoid brachiopod is from the Upper Cretaceous of
New Zealand where specimens of an undescribed species have been collected from the

KINGENINAE

beds cropping out at a stream section in the district of Koranga, North Island.
Specimens obtained by collectors in the Geological Survey of New Zealand have
been examined (locality nos. 5792, 1401) and appear in a dark-grey, highly glaucon-
itic, coarse sandstone with Ostrea lapillicota. According to Wellman (1959 : 138,
149) this fossil serves as a marker in the Lower Haumurian beds of Maastrichtian
equivalent.

Outside the Cretaceous, reference to three fossil examples of Kingena and one from
Recent seas have been made. Of these, K. raincourii was described by Deslong-
champs (1863 : 294) from the Eocene, Calcaire Grossier, of Paris basin. From the
illustrations of the specimen figured by Deslongchamps (pi. 8, figs. 12-14) fine pus-
tules or granules are seen on the shell surface but, apart from this feature, there is
little to commend its reference to the genus Kingena, In the same way, the speci-
men described and figured by Cossman & Pissaro as K, constantinensis (1904 : 31, pi.
19, figs. 29, 30) from the Eocene of the Cotontin, France, is probably not referable
to Kingena.

Likewise, the reference to Kingena simiensis by Waring (1917 : 73) from the
Eocene of Simi Hill, Ventura County, California, U.S.A. is also likely to belong to
some other genus.

A Recent form from the Indian Ocean described by Joubin (1906) as ? Kingena
alcocki has been referred by Dall (1920) and Jackson (1921) to the genus Frennlina
but it is more likely that it has closer affinities to Laqueus. An example of “ Kin-
gena” alcocki (Joubin) is figured here, PI. 12, figs. 3 & 4, for comparison with Waconella
gen. nov. with which it appears to have much in common. It is also used to illustrate
the different mode of attachment of brachial loop to median septum from that seen in
the Kingeninae.

VI. MORPHOLOGY AND STRUCTURE

The terms used in the description of the characters, both generic and specific,
throughout this work are those found in the glossary of morphological terms in the
Treatise on Invertebrate Paleontology (1965).

Special reference is made in the systematic descriptions to terms applied to charac-
ters observed from transverse serial sections representing species examined or from
dissected and acid developed specimens. Any reference to characters seen in
“ Stereoscan " micrographs or in thin longitudinal sections or transverse shell sections
has been stated in the legend to the illustrations.

External morphology. The general outline of the four genera at present repre-
sented by the subfamily Kingeninae is oval-pentagonal to pentagonal. The term
pentagonal, in this instance, being comparable to spade-like ” with the broadest end
of the spade representing the posterior part of the brachiopod.

Within this rather restricted morphological range, certain distinct features can be
recognized and are discussed in broad outline in this section, but are dealt with in
greater detail under the description of each genus. In the same way, the more subtle
details which allow the genera to be separated into species, are discussed more fully
in the systematic section.

The general morphological pattern for both Ziitelina and Kingena is very much the

THE BRACHIOPOD SUBFAMILY

Table 3

EUROPEAN

AMERICAN

STAGES

FORMATIONS

Lower

BUDA

Cenomanian

LIMESTONE

GRAYSON MARL

1 1 1

MEMBER

1—

main STREET
LIMESTONE

member

LL.

PAWPAW SHALE

MEMBER

WENO MARLY LST.

Albian

MEMBER

DENTON MARL

MEMBER

FORT WORTH

L 1 M ES TONE

DUCK CREEK

FORMATION

Kl AMICH 1

FORMATION

Correlation of the beds of the Washita Group of Texas and Mexico with the main European

stages, Albian and Cenomanian.

KINGENINAE

same. Both genera pass through a similar ontogenetic series of outlines. These
begin with subcircular, becoming broadly-oval and passing to elongate-oval to
pentagonal.

The appearance of the shell surface depends largely upon the mode of preservation,
but it is noticeable that in some species, particularly of Kingena, the concentric
growth-lines are very much more prominent.

Ornamentation of the shell surface in the Terebratellacea usually manifests itself
in the form of strong bifurcating or non-bifurcating radial costae and striae. Some
shells, however, remain free of additional ornament and have a smooth shell surface
on which can be seen the pattern of punctae. Kingena, unlike all other Terebratel-
lacean brachiopods, retains a shell surface ornament of evenly distributed spinose
pustules or granules from its earliest developmental stages to adult form. Although
this pustulate or granular ornament appears to have no parallel in either Recent or
fossil genera, comparison has been made with forms which exhibit protuberances or
outgrowths of primary or secondary shell layers.

Pustulate or spinulose external ornamentation of the shells of Productida and
Spiriferida is a common enough character in Middle and Upper Palaeozoic genera, but
it is rarely found in the Terebratulida. In the Mesozoic, however, examples of
ornament similar in some respects to that of Kingena exists on the surface of some
genera. In particular, the genus Pseudokingena, described by Bose & Schlosser
(1900) for Davidson’s Terehratula deslongchampsi from the Middle Lias of Normandy,
exhibits an apparently similar shell surface ornament to that of Kingena and is in some
ways comparable. Representative specimens of this genus are extremely rare but,
from the few examples examined, it seems possible that Pseudokingena might repre-
sent an early stage in the ontogeny of a Terebratellid rather than with the Tere-
bratulids with which it is at present classified (Muir-Wood 1965 : H 772).

In some very young stages of Recent Dallinidae an ornament of comparatively
large rounded granules has been noted. Deslongchamps (1884, 13, 14)

illustrated this ornament in a series of young stages of Macandrevia cranium (Muller).
It would appear that this type of ornament does not survive into the adult develop-
mental stages of the shell as there is no evidence of any such ornament found on
mature shells of this genus.

A '‘tear-drop” type of ornament appears on the surface of Terehratula'' ovata
(J. de C. Sowerby) and “ T. ” arcuata (Roemer) from the Cenomanian of English and
German localities. Davidson (1852, pi. 4, figs, iic & 14c) figured an enlargement of
this ornament on a specimen of “ T. ” ovata from the Cenomanian of Warminster,
Wiltshire. This type of ornament differs from that of Kingena in its shape and dis-
tribution. The “tear-drop” or pear-shaped pustules are arranged in rows, almost
equally spaced and occurring on the shell surface exposed between the numerous
prominent growth-lines as though they were originally just peripheral ornament on
the leading or growing edge of the shell.

In the longitudinal tangential sections of the shell of a specimen of Kingena
pentangulata (Woodward) from the Upper Chalk of Norfolk, shown here PI. 13, figs.
5 & 6, it will be seen that the pustulate ornament consists of a conical outgrowth of the
primary shell layer which does not influence nor is influenced by the underlying

4-2

THE BRACHIOPOD SUBFAMILY

secondary shell layers. It differs from the spines or spinules found on the surface
of some Productida and Rhynchonellida in that it is entirely composed of solid shell
matter and is not a hollow structure penetrating the shell (Williams & Rowell 1965 :

Specimens of Kingena spinulosa (Davidson & Morris) which have been examined
in preparation for this work, appear to have a somewhat distinct pustulate ornament.
Although they do not differ in structure from the pustules on the surface of other
species of Kingena, they consist of both large and small conical outgrowths and are
indiscriminately distributed over the shell surface, the smaller pustules appearing
between the larger.

Tubercles on the inner surface of the shells of some Recent Terebratellida are in
complete contrast to the structures described on the shell surface of Kingena. They
are, in some ways, more comparable to the pseudopunctae seen in the strophomenoid
shell (Williams & Rowell 1965 : H. 71) where outgrowths of secondary shell appear
on the inner surface of the valve. The tubercles of the Terebratellida, mentioned
above, are seen to have a deep underlying influence on the secondary layers in very
much the same way as do the pseudopunctae.

The umbonal features of the Kingeninae are of specific importance and may
present some evidence of change of environment. A large foramen truncating a
massive umbo is almost certainly developed as a result of the need for increased
pedicle support. Such support would be necessary in more disturbed or turbulent
waters and this character is noted more frequently on species collected from beds
exhibiting other signs of turbulence. At Wilmington, south Devon, current bedding
is a common feature of the loose sandy facies of the Cenomanian which yields numer-
ous specimens of Kingena arenosa (d' Archaic), a species with a large foramen truncat-
ing a massive umbo. The same type of umbonal characters are noted in the species
Kingena blackmorei sp. nov., PI. 14, fig. 4, from the Upper Senonian, Gonioteuthis
zone of Wiltshire and Hampshire.

A slightly produced, rounded, dome-like umbo is a character more commonly
found in the Middle and Upper Chalk species, PI. 14, figs, i & 2. Both Kingena elegans
sp. nov. from the Turonian and K. pentangulata (Woodward) from the Upper
Senonian are characterized by this strong specific feature. In the case of the Turonian
form, the foramen is small and circular and the beak-ridges indistinct or smooth. A
short interarea is developed and disjunct deltidial plates are exposed.

In Kingena pentangulata, the same dome-like umbo is developed, but there is a
strong tendency to labiation of the foramen and the interarea is extremely short,
allowing little or no exposure of the deltidial plates, PI. 14, fig. i.

Apart from the external pustulate ornament of Kingena, the general account of the
morphology of the genus, given here, might very well be applied to Zittelina from
the Upper Jurassic of Germany. The characteristic pentagonal outline prevailing
throughout the subfamily is the dominant feature of the general outline of the type
species, Zittelina orhis (Quenstedt) . Variation extends to slightly more quadrate and
subcircular plan. The genus does not show any marked tendency towards elonga-
tion, as is noted in most species of Kingena described here. Anterior sulcation of
both brachial and pedicle valves is slight and is usually bordered by faint carinae.

KINGENINAE

The umbo in Zittelina is slightly incurved, but the interarea is extensive in most
species and marked by sharp, or distinct, permesothyrid beak-ridges. The foramen
in all species remains circular and is medium to small in size.

The genus Beloihyris is represented by five species, four of which Smirnova (i960)
described from the Lower Cretaceous of the Northwest Caucasus. The fifth species,
B, pseudojurensis was originally described by Leymerie (1842) as Terebratula pseudo-
jurensis from the Neocomian of MaroUes, France, and is referred here to the genus
Beloihyris Smirnova i960.

The genus varies in outline from distinctly elongate-pentagonal to elongate-oval.
The type species B, plana Smirnova, is broad, flat and somewhat zeilleriform or
cinctiform in general outline. An anterior sulcus is fairly well marked in both
valves, giving rise to a ligate anterior commissure. The hinge-line is narrow and the
shoulders of the umbo are steep with an acute apical angle. The small circular fora-
men is bordered by permesothyrid beak-ridges.

Fig. 2. Diagram illustrating the three stages in the loop developments of Kingena and

Zittelina.

(a) Precampagiform. (b) Early campagiform.

As no specimens of the genus Tulipina Smirnova, 1962 have been obtained for the
present study, nothing can be added to the original description of the genus (Smir-
nova 1962 : 102).

Internal characters. Although the two genera Zittelina and Kingena have many
external morphological features in common, this is not to say that the two forms are
in any way more closely related than any other genera investigated here. With such
a list of characters in common, and bearing in mind their stratigraphical relationship,
it would seem logical to suppose that Zittelina was the direct ancestor to Kingena,

THE BRACHIOPOD SUBFAMILY

although investigation of the Hauterivian genus Belothyris, carried out here, throws
light on an alternative evolutionary trend and a possible link with Lower Albian
species now considered to belong to the genus Kingena. There are still many gaps
in our knowledge of these forms and more research is needed before any hard and
fast conclusions can be reached as to their lineage.

The internal characters of both Zittelina and Kingena are broadly discussed and
compared here.

Both genera have similar cardinalia, consisting of a very small cardinal process,
with deep concave hinge-trough supported by long, high median septum. The
descending branches of the brachial loop, originating from the distal ends of the
hinge-trough, extend anteriorly, developing long crural processes. Attachment of the
descending branches to the septum is by means of two broad bands, one from either
branch, which curve inwards towards the septum and then upwards and outwards.

A B

Fig. 3. A. Descending branches of Kingena showing a. points of connection to kingeni-
form hood. b. broad transverse bands to septum, c. long median septum. B. Kingeni-
form hood showing d. extensions for connection with median spetum. e. broad conical
expanded band or hood,

describing a low arc or semicircle, to form a connection for the extended ends of the
broad transverse band or conical kingeniform hood of the ascending branches.
Fig. 3, These fuse with a thickened part of the edge of the median septum at this
point.

The ascending and descending branches are broad, somewhat resembling those of
Campages, but are separated by extensive lateral lacunae. These lacunae tend to be
more highly developed in Kingena than in Zittelina,

In the Recent genera Frenulina and Laqueus which have a similar loop development
to that of Kingena and Zittelina, the brachidia are developed at a more anterior

KINGENINAE

position in the shell and appear to reach maturity at a later stage than in the fossil
genera. Kingena and Zittelina both have a mature loop developed at a very early
stage in their ontogeny.

Although lack of suitable young forms prevents a thorough investigation of the
growth stages in Zittelina, it would seem from the examination of transverse serial
sections of mature individuals that the main differences between Zittelina and Kin-
gena lie in the development of the transverse band or hood, which appears to be less
conical or more quadrate in tranverse outline in Zittelina than in Kingena, and in the
descending branches being placed at a more acute angle in relation to the central
septal pillar. In transverse serial sections of the descending branches of Kingena,
they are seen to lie almost at right-angles to the septum, except in the very early
growth-stages.

Fig. 4. Impressions of adductor and diductor muscle-scars on an internal mould of
Kingena spinulosa (Dav. & Morris) from the Cambridge Greensand, a. adductor scars,
b. impression of septum, c. adductor and diductor muscle-scars in pedicle valve, d.
umbonal cavity in pedicle valve.

In Belothyris, the median septum is almost as proportionately long as in Zittelina
and Kingena, but is higher in its development from the floor of the brachial valve.
No direct or indirect attachment is made from the septum to the descending branches.
The hood, almost triangular in transverse section, is approximately conical in general
shape and develops later or at a more anterior position in the valve than in either
Kingena or Zittelina. The general plan of the mature brachial loop, as seen in
transverse sections. Fig. 14, is to some extent reminiscent of some species of Kingena.
In particular, K. lemaniensis, which has a proportionately higher median septum than
in other species and has a somewhat similar transverse outline of the hood. The
small processes or flaps ” to which Smirnova (i960 : 117) refers in her description of
the hood of Belothyris were compared by her to the lateral flaps '' or flanges seen on
the transverse band of Gemmarcula aurea Elliott from the Upper Aptian of Faringdon,
Berkshire. Processes or flanges similar to those noted by Smirnova have been seen
on the lateral extremes of the transverse band or hood of K. lemaniensis and K.
convexiformis from the Shenley Limestone, Leighton Buzzard, Bedfordshire. It is
thought, here, however, that these processes are in no way comparable to those of

THE BRACHIOPOD SUBFAMILY

Fig. 5. Cameralucida drawing of the brachial loop of Zittelina impressula (Quenstedt)
from the Upper Jurassic, Malm, Germany. B. 45092. X 15 approx.

Fig. 6. Cameralucida drawing of the mature brachial loop of Kingena pentangulata (Wood-
ward) from the Upper Chalk, mucronata Zone, Norwich, Norfolk, showing the length of
the septum in proportion to the length of brachial valve. BB. 45789. X 20 approx.

KINGENINAE

Fig. 7. Drawing of the brachial loop of Laqueus californicus from Recent seas off California
showing a. connecting bands to septum, b. comparatively long septum, c. laqueiniform
loop. d. lateral lacunae in transverse hood. X 3 approx.

Fig. 8. Drawing of Frenulina sanguinea from Recent seas off the Philippines, a. frenulini-
form hood. b. descending branches with broad connecting bands to the short septum,
c. cardinal process, d. median septum, x 4.

THE BRACHIOPOD SUBFAMILY

Gemmarcula aurea Elliott. They are smaller in proportion to the rest of the loop and
are directed ventrally, whereas, in Belothyris plana, these structures are seen to be
deflected dorsally.

The brachial loop of Tulipina is known only from the serial sections of the type
species Terehr alula koutaisensis (de Loriol) published by Smirnova (1962). It shows
the double attachment of the loop branches to median septum but also shows that no
hood or expanded transverse band is developed, the ascending branches being
attached directly to the septum. The descending branches form an attachment to
the septum at a point when they are almost at right-angles to the septum, that is,
late in their development.

The short lateral concavo-convex plates at the base of the median septum are
unlike any structure so far examined in the Kingeninae. They may have provided
additional support to the lophophore.

In Kingena and Zittelina, as in some genera within the ZeiUeriidae, the brachial
loop is seen to have numerous small spines attached to the outer surface of the
branches. These spines originate at the extreme posterior end of the loop, becoming
more elongate anteriorly and at the distal ends of the loop. No such spines occur in
the Recent genera Laqueus and Frenulina but additional spiny outgrowths have been
observed on the dorsal surface of the branches in early loops of Macandrevia cranium,
but these disappear as the loop develops towards maturity.

Definition of relative size in the Kingeninae. For the purpose of this work
only, the term LARGE is applied to a specimen with external dimensions of approxi-
mately :

15-26 mm long,

10-16 mm wide.

MEDIUM would approximate :

10-14 mm long,

4-9 mm wide.

SMALL would be approximately :

2- 9 mm long,

3- 8 mm wide.

Variation within these limits occur in some species, but these are always noted in
descriptions.

VII. SHELL STRUCTURE

Rapid advances have recently been made in the study and understanding of the
shell structure of both articulate and inarticulate brachiopods. Such advances are
due largely to the splendid series of papers on this subject by Williams (1956, 1965,
1966, 1968) who has revived the interest sparked off by Carpenter (1843, 1855), King
(1867) and Blochmann (1908) and has projected a further dimension into the concept
of the phylogeny of the group.

The Cambridge ''Stereoscan*' electronmicroscope has been used in the present
investigation for the purpose of comparing mosaic patterns on the inner surface of
the shells of fossil species of Kingena with those of some Recent Dallinidae and other
Terebratellacean forms.

KINGENINAE

In general terms, the mosaic pattern in the Kingeninae consists of a fairly even
distribution of acute diamond shaped ends to the terminal faces of the secondary shell
fibres, not altogether different from any other Terebratellacean brachiopods. Minor
differences, however, are present which can only be explained by comparison with
Recent forms which are related, either directly to the Dallinidae or diverge to such an
extent as to allow a contrast to be drawn.

The general pattern of the shell mosaic in Kingena appears to conform to that
observed in Recent Dallinidae. The distribution and relative size of the endopunc-
tae, on the other hand, differ considerably. In K. lemaniensis shown here, PL 12,
fig. 3, the endopunctae appear large in comparison to the terminal ends of the fibres,
whereas in other Dallinidae of Recent origin, such as Macandrevia cranium, PI. 12
fig. 6, and Campages hasilanica, PL 12, fig. 4, the endopunctae are considerably
smaller and more numerous than in Kingena. They also appear to be arranged in
distinct rows almost equidistant from one another. This arrangement of the endo-
punctae seems to have more in common with Terehratella dorsata, PL 12, fig. 2, which
has a more numerous distribution of caecae.

By way of contrast, the mosaic pattern of two species of Laqueus shown on PL
13, figs. I & 2, is seen to have a more quadrate outline or less acutely diamond-shaped
than in the Kingeninae. Even at points just anterior to the muscle-marks and also
at the anterior margin the general outline of the mosaic in Laqueus is consistently
more quadrate.

VIII. SYSTEMATIC DESCRIPTIONS

Elliott (1948 : 311) erected the subfamily Kingeninae for Kingena Davidson, 1852.
The extraordinary differences in brachial loop structure, such as, the double attach-
ment of brachial branches to septum and the expanded transverse band or kingeni-
form hood in the mature Kingeninae which Elliott recognized, are here thought to be
sufficiently distinct as to warrant further separation from the main family, Dallinidae.
The family Kingenidae Elliott, 1948 is here proposed, nom. transl. for Kingeninae
Elliott, 1948.

Superfamily TEREBRATELLACEA King, 1850
Family KINGENIDAE Elliott, 1948 nom. transl.
Subfamily KINGENINAE EUiott, 1948
Genus KINGENA Davidson, 1852

Type-species. Terebratula lima Defrance, 1828

Kingia Schloenbach, 1865
Kingena Stoliczka, 1872
Kingena Dali, 1877
Kingena Hall & Clarke, 1894
Kingena Schuchert, 1897
Kingena Schuchert & LeVene, 1929
Kingena Hertlein & Grant, 1944
Kingena Elliott, 1965

THE BRACHIOPOD SUBFAMILY

Emended diagnosis. Small to large biconvex, elongate-oval to elongate-penta-
gonal ; rectimarginate to faintly uniplicate to ligate ; test thin, pustulate. Umbo
short, massive to slightly produced, suberect ; foramen subcircular, permesothyridid ;
deltidial plates disjunct. Muscle marks and pallial sinuses faint. Cardinal process
small, hinge-trough concave, shallow, broad, supported by thin long, moderately high
median septum. Loop with modified transverse band, or funnel-shaped hood
doubly attached to septum, developed from early precampagiform and campagi-
form stages.

Emended description. The genus comprises a group of variable but distinct
biconvex, pentagonal Terebratellacean brachiopods, ranging in length from less than
five millimetres to over twenty-six millimetres, and in width from three to approxi-
mately eighteen millimetres. They can be distinguished from all other Terebratel-
lacean brachiopods by the pustulate or spinulose ornament covering both valves.

Internal characters. The umbonal cavity of the pedicle valve is infilled with callus,
uniting the dental lamellae and afixing them to the floor of the valve.

Although past descriptions have always included the presence of a cardinal process,
this character is particularly poorly developed and is rarely seen in transverse serial
sections. It is composed of a flattened concave disc situated at the extreme pos-
terior end of the brachial valve.

The dorsal surface of the brachial loop is covered with short spines which project
into the brachial cavity and may have been used as additional support for the
lophophore.

The descending branches originate from the distal end of the hinge-trough and
curve in towards the septum, extending a short, broad band of connection, fusing
with the septum and connecting with the extended ends of the broad, conical trans-
verse band or kingeniform hood of the ascending branches above it.

The median septum in the brachial valve is long, extending to over two-thirds the
length of the valve.

The genus has an estimated geological range of Lower Albian to Lower Maastrichtian.

Kingena lemaniensis (Pictet & Roux)

(PI. I, figs. 1-4)

1847 Terebratula lemaniensis Pictet & Roux : 538, pi. 51, figs. 5-7.

1863 ij'erehratula) Megerlia lima Davidson ; Ooster : 35, pi. 12, fig. 16.

1900 Terebratula lemaniensis Pictet & Roux ; Ficheur : 573.

1903 Kingena newtoni Walker : 259, pi. 18, figs. 5, 7.

Emended diagnosis. Elongate-oval to elongate-pentagonal Kingena with a
distinct terebratulid aspect. Rectimarginate to markedly uniplicate. Unbo sub-
erect, short, massive. Beak-ridges distinct ; interarea extensive, deltidial plates
well exposed. Foramen large, circular.

Lectotype. CB. 2530 here selected from fifty-two specimens in the Pictet Col-
lection at the Musee d'Histoire Naturelles, Geneva, Switzerland. It is figured here,
PI. I, figs. la-c.

KINGENINAE

Horizon and locality. Lower Albian, Grand Bornand, France.

Description. The largest species of Kingena so far recorded from any locality.
It maintains an average length of approximately 25 mm., width of 20 mm., and thick-
ness of 16 mm.

As in all other species of Kingena, the early stages of growth in K, lemaniensis are
almost circular in outline becoming progressively elongate in intermediate and adult
stages.

Decidedly terebratulid in general outline, it develops a well marked uniplication of
the anterior commissure due to lateral constriction of the anterior part of the shell in
the later stages of growth. There is a fair degree of marginal thickening in the
gerontic stages.

The umbo, though massive, is slightly produced and the extensive interarea well
exposed. The permesothyridid beak-ridges which border the interarea are sharply
defined.

Although the shell surface is covered with the characteristic pustules or granules,
they are seen on the typical form from French and Swiss localities only when enough
shell has been preserved. This is a rare occurrence, but when seen, they appear as
small or fine rounded pustules, evenly distributed over the shell surface. This type
of pustulation is more often seen on the shells of the British equivalent of the species
at Leighton Buzzard, Bedfordshire. Here, however the pustules are not as well
marked on mature individuals as on the shells of young forms.

Internal characters. Essentially the same as for the genus but, from an examination
of a reconstruction made of the brachial loop from transverse serial sections, it is
possible to determine certain minor variations from the typical loop structure of
Upper Chalk species. These are seen as a less developed kingeniform hood and a
more brief attachment of descending branches to septal pillar. The serial sections
have been compared with those of Kingena pentangulata (Woodward) from the
Upper Senonian of Norfolk and figured here. Fig. 13.

Remarks. Kingena lemaniensis is a notable brachiopod species in a typical faunal
assemblage which characterizes the Lower Albian deposits of Mont Saxonet, Perte du
Rhone, Goudiniere and Reposoir in the Haute Savoie, France. Specimens from these
localities are usually found as dark-brown or black phosphatic internal moulds,
although a few specimens have been found with quite well preserved shells.

The species occurs in Britain and was described as Kingena newtoni by Walker
(1903 : 258) from the Lower Albian, Shenley Limestone of Leighton Buzzard,
Bedfordshire, Here it is found in association with a fauna of Terebratulidae and
Rhynchonellidae in every respect comparable to that found at the type locality in
France. A prominent associated species is Cyclothyris antidichotoma (Buvignier)
which was recently re-described by Owen (1962 ; 47) from the Gault of the
Ardennes, who drew attention to the similarity between specimens described and
figured by Pictet (1872 : 41, pi. 199, figs. 13-17) and those described from Leighton
Buzzard. Burrihynchia leightonensis (Walker) and Cyclothyris mirahilis (Walker)
are also found in association with K. lemaniensis. Ficheur (1900 : 573) recorded
K. lemaniensis in association with “ Rhynchonella ** cf. antidichotoma (Buvignier)

THE BRACHIOPOD SUBFAMILY

from the Lower Albian of Algeria. Ooster, (1863) refers to Terehratula lemaniensis
specimens in the Berne Museum, Switzerland, which show radiating striae on the shell
surface, a feature also noted on well preserved specimens from the British locality.

A species which was described by Walker (1903) as Zeilleria convexiformis and
which is a common fossil in the Shenley Limestone at Leighton Buzzard, may be
related to K. lemaniensis but appears to be more elongate with a shorter, truncated
umbo and steeper flanks than K, lemaniensis. The anterior commissure is recti-
marginate to very faintly uniplicate. It is referred here to Kingena and is figured
PL I, figs. 5 & 6.

Material. Numerous specimens in the Walker Collection, British Museum
(Natural History) of both K. lemaniensis and K. convexiformis. At least fifty
specimens in the collections of the Mus^e d'Histoire Naturelles, Geneva.

Range. Lower Albian.

Kingena spinulosa (Davidson & Morris)

(PI, 2, figs. 1-6, PI. 3, figs. 1-7, Text-fig. 9)

1847 Terehratula spinulosa Davidson & Morris : 253, pi. 18, figs. 6a-c.

1852 Kingena lima Defrance ; Davidson : 42, pl. 4, figs. 24 a-d.

1868 Kingena lima Defrance ; de Ranee : 163.

1869 Kingena lima (Defrance) ; Wiltshire : 185.

1874 Kingena lima Defrance ; Price : 353.

1900 Kingena lima Defrance ; Jukes-Browne : 79.

1961 Kingena sp. ; Peake & Hancock : 303.

Emended diagnosis. Elongate-pentagonal to elongate-oval Kingena, approxi-
mately 15 mm. long, 14 mm. wide and 9 mm. thick. Rectimarginate to incipiently
uniplicate. Umbo suberect, short, massive, truncated by large permesothpdid
foramen. Beak-ridges sharp. Deltidial plates, disjunct, well exposed. Interarea
extensive. Shell surface covered with large and small pustules or granules. Kingeni-
form loop supported by long median septum.

Holotype. B.5260 in the Davidson Collection, British Museum (Natural History)
and housed with three other specimens of the same species in a small box. On the
back of the box, in Davidson's handwriting, are the words Kingena lima Defr.,
Gault, Folkestone. The largest of these specimens is the original figured and
described by myself in the Annals & Mag. Nat. Hist. pl. XVIII, 1847. ”

Description. In the early stages of growth, the species is almost completely
circular in general outline, but with anterior growth, becomes pentagonal to elongate-
oval in outline in the later stages.

The lateral profile varies from plano-convex to biconvex. The characteristic
ornament of pustules or spinose granules occurs in this species as a mixture of both
large and small granules, almost conical in shape, and distributed evenly but more
sparsely over the shell surface than in other species. Enlarged photographs of the
shell ornament in K. spinulosa are shown on Pl. 13, figs. 3, 4 for comparison with
similar ornament on the surface of K. lima (Defrance), the type species from the Upper

KINGENINAE

Chalk of Margate, Kent. Davidson’s enlarged drawings of the shell surface of his
specimen (pi. 4, fig. 24d) is somewhat misleading, as it shows the tops of the pustule
with indentations or depressions which might infer that he thought them to be hol-
low, as is the case with true spinose ornament. In fact, as has been discussed
earlier, these pustules or granular outgrowths are formed from the primary layer of
the shell and are completely solid.

01 )if u \ /w\

Fig. 9. Sixteen transverse serial sections through Kingena spinulosa (Davidson & Morris) from
the Upper Albian, Red Chalk, Hunstanton, Norfolk. BB. 45790. X2.

A common feature in well preserved specimens of Kingena from Albian localities
is a faint ornament of radiating striae which appears just below the surface of the
shell. The pustulose ornament gives the appearance of following the line of stria-
tion, but this is purely an illusion. The same effect is noted on specimens where the
concentric growth-lines are prominent, the pustules then assuming a semicircular or
concentric pattern.

Internal characters. As for the genus.

Remarks. Kingena spinulosa is known only from English Upper Albian locali-
ties. Although described by some authors as being fairly common in the Gault, it is
no longer easily obtainable from the type locality at Folkestone. De Ranee (1868 :
168) in a description of the section at Copt Point, designated this species, under the

THE BRACHIOPOD SUBFAMILY

name of Kingena lima Defiance, one of the main zone fossils of his Bed II, Price
(1874 : 353) in a revision of De Ranee’s section, went even further stating '' From the
frequency of occurrence of K, lima in the zone, I am adopting the conclusion arrived
at by Mr. De Ranee and propose the name of the bed, the Zone of Kingena lima, ”
He was, of course, describing his Bed X of the Gault at Folkestone.

Kingena spinulosa also occurs as a derived fossil in the Cambridge Greensand, but
here it appears as an internal mould and is proportionately smaller than the typical
form. Nevertheless, the basic diagnostic features of the species are present in well
preserved specimens. It is possible to discern the very faint muscle marks at the
extreme posterior end of the brachial and pedicle valves of these moulds. In the
brachial valve, they are seen as small triangular areas raised slightly above the surface
of the surrounding mould and are shown reproduced here, Fig. 4, p. 45. In the pedicle
valve, the marks are less distinct and appear as elongate, broadly triangular strips.
It is also noted from these moulds that the septum is proportionately longer in the
young specimens.

In the Upper Albian, Red Chalk, at Hunstanton, Norfolk, the species is also
recognized. Numerous specimens have been collected from these beds over the
past twenty or thirty years by Mr. H. Le Strange. From the specimens collected,
are well known marked variants which appear to be less elongate than those found
among the typical form. They tend towards greater width, less produced or trun-
cated umbo and some exhibit a faint sulcation of the anterior part of the brachial
valve. The early stages throughout the full range of variation, however, agree in
every detail with those found in the Gault at Folkestone and from the derived fossils
in the Cambridge Greensand.

Although the geographical distribution of this species probably extends as far as
the French Upper Albian, no records from continental localities can be found in the
literature.

Muir-Wood (1930 : 34) described and figured a specimen under the name of Kingena
spinulosa (Davidson & Morris) from the Gault of the Samana Range, India. The
very poor internal moulds with which she illustrated the species are considered here
to be in no way comparable to those of Kingena spinulosa as originally defined by
Davidson and Morris (1847 • ^53, pi. 18, figs. 6, a-c). This form has, therefore, been
renamed here in honour of the late Dr. H. M. Muir- Wood as Kingena muirwoodae
nom. nov. pro. Kingena spinulosa (Davidson & Morris) ; Muir-Wood (1930 : 34,
pi. 5, fig. 6) non Kingena spinulosa (Davidson & Morris) 1847.

Range. Upper Albian.

Dimensions of topotype and other specimens.

All the above specimens are in the British Museum (Nat. Hist.) and were collected
from the Gault at Folkestone.

BB. 3296
BB. 3297
BB. 4628
BB. 4629

Length
16 mm
12 mm
II. 5 mm
9.8 mm

Breadth
14 mm
10 mm
10 mm
10 mm

Thickness
10.5 mm
7.7 mm
7 mm
6 mm

KINGENINAE

The following specimens are in the Sedgwick Museum, Cambridge, and are from the
Upper Albian, Hunstanton.

B. 41751
B. 41739
B. 41762

Length
16.5 mm
16 mm
14 mm

Breadth

15 mm

16 mm
14 mm

Thickness
10 mm
9 mm
8 mm

Kingena arenosa (d’Archaic)

(PI. 4, figs. 1-7, PI. 5, figs. 1-5. Text-fig. 10)

1846 Terehratula arenosa d’ Archaic : 324, pi. 21, figs. la-e, 2a, 3a, b.

1847 Terehratula lima d'Orbigny ; d’Orbigny : 98, pi. 512, figs. 1-5.

1852 Kingena lima Defrance ; Davidson, pi. 4, figs. 21, 22, 25.

1903 Kingena lima Defrance ; Jukes-Browne & Hill : 113, 248, 257.

Emended diagnosis. Medium to large Kingena, Broad, oval-pentagonal,
width almost equal to length. Short, massive, suberect umbo truncated by large,
circular foramen. Well defined, extensive interarea bordered by sharp beak-ridges.
Deltidial plates disjunct, well exposed. Rectimarginate to sulcate to faintly uni-
plicate anterior commissure. Growth-marks distinct. Pustulation fine, evenly
distributed.

Neotype. The d' Archaic Collection was composed largely of material borrowed
from other collectors. The whereabouts of the brachiopods used to illustrate his
monograph (1846) is unknown and most of the specimens are thought to have been
lost. No trace of the specimens described as Terehratula arenosa and figured on his
pi. 21 can be found in France or Belgium and it is therefore considered necessary to
propose a neotype for this species in order that it should once again become estab-
lished in the literature.

The specimen here proposed as neotype is from the Tourtia of Tournai, Belgium
and is in the collections of the Institut royal des Sciences naturelles de Belgique,
Brussels no. I.G. 8261. It is figured here, pi. 4, fig. 3a-c.

Description. Perhaps one of the most variable of any of the species of Kingena,
the typical form, as defined here, is nonetheless distinctive, being fairly large,
flattened form in maturity, with well marked sulcus on the anterior part of the pedicle
valve. When further developed in gerontic specimens, this sulcus gives rise to a
faint uniplication of the anterior commissure. Well developed, almost step-like
growth-lines are common features on well preserved specimens.

Unlike the ornament of Kingena spinulosa from the Upper Albian, the pustulation
on the shell surface of K, arenosa is of uniform size, being fairly fine and evenly
distributed over the surface of the valves.

A broad hinge-line in the typical form gives rise to an extensive interarea bounded
by sharp beak-ridges. The large, circular foramen truncates a short, massive umbo.
The anterior profile is elliptical.

In early growth-stages the species is plano-convex to just biconvex, becoming
acutely biconvex with, sometimes, marked inflation of the brachial umbo, which
diminishes with lateral and anterior growth of the shell. The pedicle valve maintains
its early acute convexity throughout all growth stages.

THE BRACHIOPOD SUBFAMILY

Internal characters. As for the genus. See Fig. 12, p. 61.

Remarks. Most of the specimens examined from the type locality are in museum
collections. In the main, this material has been collected from two or three sites
within the type area of Gussignies near Tournai, Belgium. The deposits at these
localities, known as the Tourtia, are generally thought to be of condensed Upper
Albian and Cenomanian age. Apart from the presence of Terehratula capillata
d' Archaic, which is known only from Albian localities in Britain, the brachiopod fauna
in the Tourtia of Belgium contains very little of Albian significance. However, until
more faunal evidence is available, the exact age of the deposits remains indefinite.

Fig. 10. Twelve transverse serial sections through Kingena arenosa (d'Archiac) from the
Tourtia of Tournai, Belgium. BB. 45784. X2.

In the same way, the '' Greensand deposits at Warminster, Wiltshire, are of
indefinite age, no correlation with coastal sections having been made so far. Here
again, the beds yield what is considered to be a typical Cenomanian brachiopod
fauna and Kingena arenosa (d'Archaic) occurs in these beds with Cyclothyris difformis
(Val. in Lamarck), '' Terehratula ovata (J. de C. Sowerby), Deretapectita (J. Sowerby
and Orhirhynchia mantelliana (J. de C. Sowerby). The Warminster beds are no
longer exposed.

In Britain, K. arenosa occurs in the sandy facies of the Middle Cenomanian at
Wilmington, Devonshire, in the Cenomanian Beds B, at Bovey Lane sandpit and
from beds of equivalent age in the coastal sections from Branscombe, Hooken,
Seaton, south Devon and along the Dorset coast to Lulworth. It is also found in the
middle Cenomanian deposits at Culver and Compton Bay, Isle of Wight.

Similarly, in France, Kingena arenosa occurs in the Lower and Middle Cenomanian
Limestone at BlevUle, and at Cap le Heve near Le Hivre, Normandy.

At Essen in North Germany, the species occurs in the Lower Cenomanian deposits

KINGENINAE

known as the “ Essen Tourtia or “ Essen Greensand '' and may be the same species
described as Terehr alula pectoralis by Roemer (1840 : 42) and figured (pi. 7, fig.
iga-c), but the illustration is poor and the whereabouts of the specimen unknown and
it is here regarded as nomen. duhium. Reuss (1845) described a specimen under the
name Terehr alula pecloralis Roemer and figured a specimen (pi. 26, fig. 12) from the
Planerkalk of Kutschlin, Czechoslovakia. This may belong to the species K.
arenosa (d' Archaic) although the specimen figured is lacking in some of the more
diagnostic features, such as, pustulation of the shell and no median septum is visible
from the figure.

Apart from the type locality at Tournai, the species also occurs at Montignies-sur-
Roc and Chercq (Pont-a-Rieu) in Belgium.

K. arenosa differs from K, spinulosa from the Upper Albian in its regular or uniform
size of pustules on the surface of the shell and in its different dimensions. It is
proportionately wider than any species of Kingena described here and the umbonal
characters are dominated by a large foramen.

Dimensions of neotype and other specimens

Length Breadth Thickness

I.G. 8261 Neotype 16 mm 16 mm 11.9 mm

Additional material
B.M. = British Mus. (Nat. Hist.)

I.G. = Inst. roy. Sci. Brussels

B.M. 88760

Length

Breadth

Thickness

Warminster
B.M. B.35456,

28 mm

29 mm

19.9 mm

Montignies-sur-Roc.
I.G. Unregd. Duplicate,

14 mm

13.6 mm

II. 5 mm

Bleville, France.

14 mm

10 mm

Further material used. B.M. B. 82713-17, Warminster. B.M. B.491 War-
minster. B.M. B. 8268-70, Warminster. B.M. BB. 9319, Lulworth Cove, Dorset.
B.M. BB. 7053-56, Bindon Steps-slips, Seaton, Devon. B.M. BB. 42412, Cran
d'Escalles, nr. Cap Blanc Nez. B.M. B. 35197, Le Havre, Normandy. B.M. B.
15474, Essen, Germany. B.M. B. 35692, Essen, Germany. B.M. B. 85103-6,
Montignies-sur-Roc, Belgium. B.M. B. 35456, Montignies-sur-Roc, Belgium. I.G.
5096, Tournia. I.G. 12515, Chercq (Pont-a-Rieu), Belgium.

Kingena concinna sp. nov.

(PI. 5, figs. 6, 7, 8. PI. 6, figs. 4-6, Text-fig. ii)

1852 Kingena lima Defiance ; Davidson : 44, pi. 5, figs, i, 2, 4.

1903 Kingena lima Defiance ; Jukes-Browne : 43, 192, 204, 224.

Diagnosis. Shell oval, acutely biconvex. Umbo massive, incurved ; foramen
small. Anterior commissure rectimarginate to just ligate. Finely pustulate
Hingeline narrow, beak-ridges short, indistinct.

THE BRACHIOPOD SUBFAMILY

Holotype. BB. 93758 in the British Museum (Natural History), collected from
the Middle Cenomanian, rhotomagense zone, (Bed 5 of Price), Foreshore between
Folkestone and Dover, Kent.

Dimensions of holotype. 15 mm. long, 13 mm. wide and 10 mm. in thickness.

Description. A small globose Kingena with a distinctly terebratuUd aspect.
The general outline is unmistakably oval but some variants have a tendency to
straighter or subparallel sides and foreshortened anterior. This variant is more com-
monly found in assemblages from the Cambridge Greensand, although it does occur
at Folkestone and Dover. The rounded umbo is slightly incurved, obscuring the
deltidial plates. The lateral profile shows the acute binconvexity of the shell and a
degree of posterior inflation of the brachial umbo.

Fig. II. Fourteen transverse serial sections through a specimen of Kingena concinna sp.
nov. from the Middle Cenomanian, rhotomagense Zone of Folkestone, Kent. BB. 45785.
X3-

Remarks. This small species has often been misidentified as Terehratula squa-
mosa Mantell and occurs with this species at Dover and Folkestone.

It is thought that K, concinna occurs in the Totternhoe Stone and possibly in the
basal beds of the Plenus Marls, but specimens collected from these beds are often
damaged or crushed. In Lincolnshire it occurs in the H, subglobosus zone (York-
shire equivalent) at Louth and has been collected from a similar horizon at Kipling-
cotes Valley, East Yorkshire by Mr. C. W. Wright.

The species can be distinguished from other described species of Kingena in its
extreme convexity or semiglobose outline and general terebratulid appearance.

Paratypes. All from the collections in the British Museum (Natural History).

B. 25030, Burwell, Cambridge. B. 51479, West Cliffs, Dover. BB. 7446, Tinhead,
near Bratton, Wiltshire. B. 55092 Pit 2, near waterworks, Hubbard's Valley,
Louth, Lincolnshire. BB, 45811, Rifle Butts Pit, Kiplingcotes Valley, E. Yorkshire.

KINGENINAE

Kingena elegans sp. nov.

(PI 6, figs. 1-3)

1968 Terebratula (Megerleia) lima Defrance ; Schloenbach ; 32, pi. 3, figs, i, 2.

1904 Kingena lima Defrance ; Jukes-Browne ; 289, 229.

1942 Kingena lima (Defrance) ; C. W. & E. V. Wright : 117.

Diagnosis. Small to medium Kingena ; elongate-pentagonal. Umbo slightly
produced. Hinge-line broad, interarea extensive. Foramen large, beak-ridges
sharp, permesothyridid. Anterior commissure sulcate.

Holotype. B. 55241 in the British Museum (Natural History), from the Turon-
ian, Holaster planus Zone, Pit 8, Boswell Farm, 3^ miles north west of Louth,
Lincolnshire.

Description. A trim, neat-looking, regular pentagonal shell with well marked
growth-lines. The average length is 14 mm., width ii mm., and thickness 9 mm.
The umbo is characteristic of the species being dome-like, suberect and with wide
interarea and well marked beak-ridges. There is an even degree of convexity of
each valve with a shallow sulcation of the anterior part of the brachial valve. A
corresponding flattening of the pedicle valve in the anterior region is bordered by
faint, slightly radiating, carinae, one on either side of the flattened area.

Internal characters. As for the genus.

Remarks. Kingena elegans was originally described as Terebratula {Megerleia)
lima from the Turonian, Galeriten-Schichten of north Germany by Schloenbach
(1868). It is the least variable of all species of Kingena and maintains its character-
istic elongate-pentagonal outline throughout the full range of its geographical dis-
tribution.

In England the species occurs chiefly in the Holaster planus zone at Kiplingcotes,
Yorkshire and at the same horizon from Louth, Lincolnshire, but specimens have
been collected from the Terehratulina lata Zone of Swaffham, Norfolk. A single
specimen, somewhat larger than the typical, has been collected by Mr. Christopher
Wood, Institute of Geological Sciences, from the Turonian, H. planus Zone of Reed
Quarry, near Royston, Hertfordshire, and probably marks the approximate southern
limits of the species in this country.

Kingena elegans differs from K. pentangulata (Woodward) in its proportionately
smaller dimensions, regular pentagonal outline, marked growth-lines and anterior
sulcation of the brachial valve.

Specimens collected from Yorkshire and Lincolnshire are often pink in colour.
This colouration of the shell is not an uncommon feature among specimens collected
from other horizons within the Middle and Upper Chalk. It is probably the remains
of natural pigment of the shell, a common enough character in some of the Recent
terebratellacean brachiopods. It is also noted on specimens of Kingena mesembrina
(Etheridge) from the Gingin Chalk of Western Australia and Elliott (1952 : 5) drew
attention to this in his description of the species.

Range. The species appears to be confined to the Turonian.

Dimensions of holotype. 11*5 mm. long, 10 mm. wide, 7 mm. thick.

THE BRACHIOPOD SUBFAMILY

Paratypes. All in the British Museum (Natural History). B. 55242-45, Boswell
Farm, near Louth. (Probably T, lata zone). B. 55222-26, Wyham House, 6 miles
N.N.W. of Louth. B. 93438-46, No. 23a. Swaffham Cutting, Norfolk. (T. lata zone).

Kingena mesembrina (Etheridge)

(PI. 6, fig. 9 a-c)

1913 Magas mesemhrinus Etheridge fit. : 15, pi. 2, fig. 5-8.

1952 Kingena mesembrina (Etheridge) ; Elliott ; 4, pi. i, figs. 1-13.

As this species has been reviewed as recently as 1952 and has been adequately
described by Dr. G. F. Elliott, it is not considered necessary to re-describe it in
detail for the purpose of the present work. Elliott’s diagnosis and a shortened
version of his excellent description are given below. A duplicate specimen obtained
from the Gingin Chalk and kindly presented to the British Museum (Natural History)
by Dr. Pat Coleman of the Geological Department, University of Western Australia,
is figured here, PI. 6, fig. 8a-c. This specimen compares favourably with the photo-
graph of the original type specimen which Etheridge figured (1913 : pi. 2, figs. 5, 6).

Diagnosis. Shell about 18 mm. long, 16 mm. wide and 9 mm. thick, biconvex
rounded pentagonal to elongate-oval, test thin, externally finely granulose and con-
spicuously punctuate, anterior commissure feebly sulcate. Umbo short suberect,
truncated ; foramen moderately large, circular, permesothyrid ; deltidal plates dis-
junct and obscured by the beak-ridges, hinge-line terebratulid. Hinge-teeth sup-
ported by dental lamellae, joined by sessile pedicle-collar. Cardinalia showing strong
fused inner socket-ridges and crural bases, wide hinge-trough and transverse cardinal
process ; kingeniform loop supported by a short median septum. Muscle marks
and pallial sinuses faint.

Neotype. The original specimen figured by Etheridge (1913) is mislaid or lost
and the whereabouts of any paratypic material unknown. A specimen figured by
Elliott (1952) is here proposed as neotype in place of the specimen which Etheridge
figured (pi. 2, fig. 5-8). Elliott’s specimen is one of twenty-one hypotypes in the
collections of the Department of Geology, University of Western Australia and is the
specimen figured by him (1952 : pi. i, figs. 12, 13).

Description. This shell varies somewhat in outline, though easily recognizable.
Specimens which are rounded pentagonal in outline include the largest individuals.
The test for the species as a whole shows on well-preserved specimens the fine
external surface granules characteristic of the genus ; the punctation is coarse and
close-set.

Remarks. The species described by Elliott seems to have a shorter septum than
in some of the European forms of equivalent age, but the rest of the internal charac-
ters appear to be comparable.

KINGENINAE

Kingena lima (Defiance)

(PI. 6 , figs, 7-8, PI, 7, fig, 4a-c, 5a-c, Text-fig. 12)

1828 Terebratula lima Defiance : 156.

1863 Terebratula (Kingena) sexradiata (J. de C. Sowerby) ; Deslongchamps : 292, pi. 7, fig. 5-8.
1900 Kingena lima Defiance ; Rowe : 300, 363.

1904 Kingena lima Defiance ; Jukes-Biowne : 42, 80, 221.

1908 Kingena lima Defiance ; Rowe : 338.

Diagnosis. Small rounded-pentagonal Kingena ; plano-convex to biconvex
rectimarginate. Umbo massive, suberect ; beak-ridges distinct. Foramen large
circular, permesothyridid. Growth-lines numerous, distinct. Pustulation uniform
evenly distributed.

Fig. 12. Ten tiansveise serial sections thiough a specimen of Kingena lima (Defiance) fiom the
Uppei Chalk, marsupites Zone, Maigate, Kent. BB. 45786. x 3,

Neotype. There is very little evidence of a type specimen or specimens having
been used by Defiance in his original description of the species in 1828. It is more
likely that, following the practice of palaeontologists of the time, he established the
species on knowledge obtained from several examples of what he considered to be a
typical or characteristic fossil of the Chalk. His locality details were vague and he
quotes “ collected from the Chalk in the district of Beauvais and the stratigraphical
details are for pure speculation.

Since there can be no certainty of either the morphological details or the strati-
graphical age of the species described by Defiance and, since the Defiance Collection
was, in any case, destroyed by the allies during the second World War, a neotype
for Kingena lima (Defiance) is proposed here.

Difficulty in obtaining suitable material from any French localities within the

THE BRACHIOPOD SUBFAMILY

Beauvais district or any comparable site, has led me to select a specimen from the
Upper Chalk of Marsupites testudinarius Zone from Foreness Point, Margate, Kent
and is in the collections of the British Museum (Natural History) no. B, 79709. It is
figured here, PI. 7, figs. 4a-c.

Dimensions of neotype. 12 mm. long, 11.4 mm. wide and 6 mm. thick.

Description. The length and breadth of this species are almost equal, but vari-
ation extends to more elongate or elongate-oval and more robust or more acute
convexity. A very strong feature of the umbo is the proportionately large foramen,
sharp beak-ridges and marked extensive interarea exposing the conjunct deltidial
plates.

The anterior commissure remains rectimarginate but a faint sulcus is developed
in the margin of the more elongate variants or older individuals.

K. lima can be distinguished from aU other species of Kingena by its very charac-
teristic rounded-pentagonal outline, plano-convex profile and marked umbonal
features. It is one of the smallest species described.

Because of the comparative rarity of occurrence of this species, records of distribu-
tion and horizontal range have been obtained largely from collected material found
in museum collections. It appears, however, to range from the Micr aster cortes-
tudinarium zone of the Lower Senonian to the Marsupites testudinarius zone of the
Middle Senonian. It is more commonly found in the Uintacrinus socialis and
Marsupites bands in the Senonian at Margate and St. Margaret’s Bay, Kent. It is
also fairly common at equivalent horizons in the Chalk of Angmering and Burpham
in Sussex. A few specimens, although somewhat fragmentary, have been obtained
from these bands at Flamborough Head, Yorkshire.

Material. M. cortestudinarium Zone. B. 79797-8, Dover, Kent. B. 93483,
Swaffham, Norfolk.

M. coranguinum Zone. B. 79691-705, Thanet Coast. B. 79706, St. Margaret’s
Bay, Kent.

Uintacrinus socialis Zone. B. 97084, Devizes Rd., W. of Salisbury. B. 79710-23,
Thanet Coast. B. 91929-41, Highfield, W. of Salisbury.

Marsupites testudinarius Zone. B. 92049, Bishop Down, nr. Salisbury. B. 47968,
Pit. 29, Highdown Hill, E. of Angmering, Sussex. B. 47974, Pit. 27, Burpham river
cliff, S. of St. Mary’s Church, Burpham, Sussex. B. 96216-7 Foreness Point, Mar-
gate. Kent.

All the above specimens are in the collections at the British Museum (Natural
History) .

Kingena sexradiata (J. de C. Sowerby)

1850 Terehratula sexradiata J de C. Sowerby in Dixon : 346, 354, pi. 27, fig. 10.

1852 Kingena lima (Defrance) ; Davidson : 42.

Emended description. Small Kingena measuring 8*9 mm. in length and 8*5 mm.
in width. The measurement of the thickness cannot be estimated owing to the
nature of the preservation of the specimen. The specimen is slightly crushed and

KINGENINAE

completely surrounded by the chalk matrix on the ventral side. It is possible to discern
a faint, shallow sulcus on the brachial valve which originates from the umbonal
area and broadens gently anteriorly. This sulcus is bordered by faint carinae, one
on either side. The very faint radiating striae which Sowerby observed are just
visible, but appear to number more than six. The median septum is clearly visible
and extends to over two-thirds the length of the brachial valve. The shell surface is
finely pustulate, the pustules being of even size and distribution. A faint pink
colouration of the shell is noticeable.

Holotype. In the British Museum (Natural History), no. 30584, from the Chalk
of Sussex.

Remarks. This is a very doubtful species since it is not known from exactly
what horizon the original specimen was collected. The type specimen is the only
specimen known to exist.

Kingena blackmorei sp. nov.

(PL 7, figs. 1-3, PI. 8, fig. i)

1904 Kingena lima Defrance ; J ukes-Bro wne ; 66, 85.

1964 Kingena pentangulata (Woodward) ; Muir-Wood : 148, pi. 52, fig. 6.

Diagnosis. Kingena, broadly pentagonal, width often equal to length. Plano-
convex to biconvex. Rectimarginate to incipiently uniplicate. Hinge-line broad ;
interarea extensive, deltidial plates well exposed. Umbo short, massive, truncated.
Foramen large, subcircular. Median septum long. Pustulate ornament fine,
evenly distributed. Growth marks numerous, indistinct.

Holotype. B. 92779 in British Museum (Natural History), collected from the
Upper Senonian, Gonioteuthis quadrata Zone, East Harnham, Wiltshire and figured
here, PI. 7, fig. 2.

Description. A large Kingena with an average length of 18 mm., width 17*6
mm. and thickness 10-4 mm. Although characteristically subcircular in early
growth-stages, it remains constantly pentagonal through later stage of growth and
in gerontic development. The brachial valve remains flattened in all stages,
developing a very slight convexity in the umbonal region in early growth and main-
taining a slight inflation of this part of the valve in later stages. The length of the
median septum in the brachial valve, though somewhat variable, is always over two-
thirds the length of the shell. In some individuals the septum has been seen to
extend to within 2 mm. of the anterior margin in the brachial valve.

A large subcircular foramen dominates the pedicle umbo, truncating it. Permeso-
thyridid beak-ridges are prominent and border a well marked, extensive interarea.

Internal characters. As for the genus.

Remarks. This species characterizes the chalk of the Gonioteuthis quadrata Zone
in the Salisbury district of South Wiltshire and also occurs in the Upper Chalk beds
of the same horizon in Hampshire, a fine series of specimens having been collected
from excavations for the Southampton Waterworks.

THE BRACHIOPOD SUBFAMILY

It also occurs in Belgium and has been collected from the Craie de Trivieres at
Harmignies and Obourg, Specimens have also been found in the Conglomerat
d’Obourg and in the base of the Craie d'Obourg,

K, hlackmorei differs from K, pentagulata (Woodward) in its less acutely convex
valves, larger foramen, less produced umbo and more prominent or sharper beak-
ridges. It also has a less constricted anterior portion of the shell. In addition,
the extensive interarea allows the exposure of the deltidial plates which, in the case
of K, pentangulata, are usually hidden by a more incurved umbo.

Unlike other species of Kingena from the Upper Chalk, the maximum width of the
shell in K. hlackmorei occurs at the posterior end of the shell, giving rise to a more
extensive or wider hinge-line.

Dimensions of holotype. 22 mm. long, 21 mm. wide and 13 mm. thick.

Dimensions of paratypes in the Sedgwick Museum Cambridge.

Length

Breadth

Thickness

B. 65638

19 mm

II mm

B. 65639

21 mm

12 mm

B. 65640

17 mm

16 mm

10 mm

B. 65641

19 mm

18 mm

10.4 mm

B. 65642

18.5 mm

17.4 mm

9.9 mm

P- 65643

17 mm

17.5 mm

9.6 mm

Other material used. GSM. 10694 (Institute of Geological Sciences). IG
6292 (Inst. roy. Sci. Belg. Brussels).

Kingena pentangulata (Woodward)

(PI. 7 figs. 6, 7. PI. 8, figs. 2-6, Text-fig. 13)

1833 Terehratula pentangulata Woodward : 49, pi. 6, fig. 10.

1836 Terehratula ventroplana Roemer : 51, pi. 2, fig. 7.

1847 Terehratula hehertiana d’Orbigny : 108, pi. 514, figs. 5-10.

1852 Kingena lima Defrance ; Davidson : 42, pi. 4, figs. 16, 18, 20.

1901 Kingena lima (Defrance) ; Rowe : 64

1904 Kingena lima (Defrance) ; Jukes-Browne : 267, 482.

1908 Kingena lima (Defrance) ; Rowe : 252.

1961 Kingena lima (Defrance) ; Peake & Hancock : 320.

Diagnosis. Large Kingena, approximately i6-8 mm. long, 14-4 mm. wide and
9*3 mm. thick. Oval to pentangulate or pentagonal ; plano-convex to biconvex.
Rectimarginate to uniplicate. Evenly pustulate. Beak slightly produced, erect to
slightly incurved ; beak-ridges poorly defined. Foramen permesothyridid, circular,
medium to small. Deltidial plates obscured by incurved beak. Kingeniform loop
supported by long median septum.

Neotype. The specimen figured by Woodward (1835 ; pi. 6, fig. 10) from Harford
Bridge, Norfolk has been lost. A neotype is therefore proposed from the collections
of the Castle Museum, Norwich, registered no. 2060. This is the specimen figured
by Davidson (1852, pi. 4, fig. 20) and is part of the Fitch Collection. It is figured
here on PL 8, fig. 5. Woodward's specimen was collected from one of two pits

KINGENINAE

working the Upper Chalk at Harford Bridge (Grid. Ref. TG/221057). According to
Peake & Hancock (1961 : 336) this locality exposes the top of the Eaton Chalk and
part of the Weybourne Chalk, both high in the Senonian and within the broad
Belemnitella mucronata Zone, Upper Campanian.

In the explanation of Woodward's plate 6, he attributed T erebr alula pentangulata
to Phillips, but there is no evidence in literature that this name was ever used by
Phillips for a Terebratulid brachiopod.

Type locality. The locality given for the proposed neotype is Norwich. Al-
though this information might seem somewhat vague, the species is typical of speci-
mens not uncommonly found in the lower part of the B. mucronata Zone of the
Norwich districts.

Fig. 13. Serial sections through Kingena peniagulata (Woodward) from the Upper Chalk,
wzwcyowa/a Zone, Norwich, Norfolk. BB. 45787. X2.

Description. Young specimens of this species are difficult to recognize and are
seldom collected from the Upper Chalk. The ontogeny of the species can be traced,
however, by studying the growth-lines on mature, young and intermediate forms.
In the early stages of growth, the general outline is subcircular becoming more
elongate-oval to pentangulate or pentagonal in maturity. There is a tendency to
marginal gerontic thickening and very slight uniplication of the anterior com-
missure. The maximum width of the species occurs at about half the distance from
the umbo to anterior margin, unlike K. blackmorei sp. nov. where the maximum
width occurs at less than one third the distance from the umbo to anterior margin.

The shell surface is covered with the characteristic pustules of even size and
distribution. The punctae are small but fairly widely distributed.

0-3

0-4

THE BRACHIOPOD SUBFAMILY

Internal characters. These are essentially as described for the genus with the
exception that, as in all Upper Chalk forms, the kingeniform hood is much more
developed anteriorly.

Distribution. Records of Kingena pentangulata (Woodward) have proved
difficult to trace since Davidson (1852 : 42) referred all species of Kingena to K. lima
(Defrance) and references to the species by Rowe (1900, 1901, 1908), Jukes-Browne
(1904) and latterly Peake & Hancock (1961 : 320) have been made under the name of
Kingena lima. However, several fine examples have been examined from the B.
mucronaia Zone, near Salisbury, and two near perfect specimens from the same
horizon are in the Sedgwick Museum, B. 5454, B. 5472 and are said to have been col-
lected from the Isle of Wight.

Similarly, continental references have also been to Kingena lima (Defrance) and the
absence of well documented collections has made the work of defining the distribution
of K. pentangulata outside British localities very difficult.

In Belgium, the species is not uncommonly found in the Craie d'Obourg at Har-
mignies, Obourg and Cuesmes and the Craie de Nouvelles at Harmignies, Heure-le-
Romain and Ciply. It also occurs in the Craie de Spiennes at Harmignies and
Ciply. Here it appears as the typical pentangulate-oval form with the same degree
of variation as the Norfolk form. It may also be the species selected and listed by
Bosquet (i860 : 390) as Megerlea pusiulosa from the Upper Chalk of Limburg, but as
there is neither description nor illustration upholding this citation it must remain
nomen duhium.

In France, the species was described by d'Orbigny (1847 : 108, pi. 514, figs. 5-10)
as Terebratula hehertiana from the Senonian of Chavot and Cezane near d’Ay in the
Marne.

Remarks. Rowe (1901 : 64) considered the genus Kingena to be represented by
one species only within the Upper Chalk, namely K. lima (Defrance), but acknow-
ledged the characteristic shape variations within zonal limits, thus, in describing the
B. mucronata Zone of Dorset (1901 : 24) states Kingena lima here assumes a shape
variation which is fully diagnostic of the zone. It is large and pentangulate. These
characters begin to appear in the Actinocamax quadratus Zone and are notably seen
in Dr. Blackmore’s series from the Salisbury Chalk . . . '' and later ** This shape
variation is well figured as Terebratula pentangulata by Woodward in the * Geology of
Norfolk * ....** In pointing out these characteristic shape variations, Rowe
merely emphasizes the subtle differences which separate the forms from the Gonio-
teuihis quadrata Zone, here described as K. blackmorei sp. nov., from the species K.
pentagulata (Woodward). These differences are chiefly confined to beak characters
and position of the maximum width of the shell.

Although the length of the median septum in the brachial valve varies from just
over half to two-thirds the total length of the shell in this species, it is noticeable that,
in the more pentangulate forms where the general outline shows less anterior con-
striction, the septum is often very much longer, extending to weU over two-thirds of
the shell length, as in K. blackmorei. It is not certain whether this variation in the
length of the septum could have any stratigraphical significance. More accurately

KINGENINAE

collected material at all growth stages must be examined before any suggestion of
possible stratigraphical value in this character can be seriously considered.

The ovahpentagonal outline of this species is recognizable as one of the early
ontogenetic stages which is maintained in later growth and at all stages throughout
its comparatively short stratigraphical range. Variation tends towards a more
predominantly oval outline in forms from the higher zones of the Upper Senonian and
early Maastrichtian.

Recent work on the stratigraphy of the Cretaceous of Northern Ireland by Mr.
Christopher Wood, Institute of Geological Sciences, has added greatly to the records
of this species from the Lower and Upper Campanian of County Antrim. Several
well preserved typical specimens have been collected from beds equivalent to the basal
mucYonata Zone of Norfolk, as defined by Peake & Hancock (1961), and two speci-
mens, though somewhat atypical in their general outline, have been collected from
beds equivalent to those within the Beeston Chalk of Norfolk, Table i.

Typical examples of K, pentangulata have been collected from the core of a boring
put down at Port More by the Institute of Geological Sciences. These occurred at
depths between 41 1 and 500 feet and all appeared to be proportionately smaller than
examples examined from English localities.

Apart from the localities in Norfolk and Northern Ireland already mentioned, the
species has been collected from Studland Bay, Dorset, Coombe Pit and Wool, south
Dorset, Alderbury, nr. Salisbury.

Kingena asperulina Stoliczka
1872 Kingena asperulina Stoliczka : 28, pi. 7, fig. 14.

Emended description. A large Kingena ^ almost circular in general outline with
a slightly produced, suberect umbo. The pedicle foramen is large and circular with
weU defined permesothyridid beak-ridges bordering an extensive interarea. A slight
sulcation develops at the extreme anterior part of the pedicle valve giving rise to
faint uniplication. Faint growth-lines and fine pustulate ornament cover the surface
of both valves.

Holotype. G.S.I. no. 1594, Geological Survey of India, Calcutta. Length,
29-50 mm., width, 27-70 mm., thickness, 20-00 mm.

Horizon and locality. Utatur group, north-west Coodycaud, Southern India.

Remarks. In general outline and size, this species closely resembles the form
from the Upper Albian, Red Chalk of Hunstanton, Norfolk, described here as
Kingena spinulosa (Davidson & Morris). It differs from that species, however, in its
slightly more produced beak, more extensive interarea and more evenly convex
valves. Since this is the only specimen mentioned and figured by Stoliczka in his
original description of the species, it is assumed that there is no paratype material
available.

The Utatur group includes beds of both Upper Albian and Cenomanian age, accord-
ing to the Lexique Stratigraphique International. The Cenomanian beds are appar-

THE BRACHIOPOD SUBFAMILY

ently commonly exposed in the Coodycaud district but it is not known for certain
whether or not the Upper Albian has ever been recorded from this area.

An approximate correlation with European beds of the Cretaceous is given here.
Table 2.

Kingena shalanurensis Stoliczka
1872 Kingena shalanurensis Stoliczka : 29, pi. 7, figs. 15-17.

Emended description. Distinctly elongate-oval in general outline with even
biconvexity of the valves. The short, slightly produced, suberect umbo has fairly
distinct but rounded permesothyridid beak-ridges and is dominated by a circular
foramen. The hinge-line is narrow, giving the umbo a somewhat terebratulid
aspect. A faint sulcus in the anterior part of the pedicle valve gives rise to incipient
uniplication of the anterior commissure.

Lectotype. G.S.T. no. 1595, Geological Survey of India, Calcutta, here selected
from three syntypes all figured by Stoliczka (1872). The lectotype is the specimen
figured by StoHczka pi. 7, fig. 15.

Dimensions of lectotype. Length, 25*50 mm., width, 24*60 mm., thickness,
14*00 mm.

Horizon & locality. ? Trichinopoly group. Possibly Lower Senonian, Shalanur,
Southern India.

Remarks. The exact age of this species must remain somewhat uncertain since
it is not known from exactly what part of the succession the specimens figured by
Stoliczka were collected. According to information taken from the Lexique Strati-
graphique International and other sources [Dr. M. K. Howarth, personal communica-
tion] the Trichinopoly group embraces beds of Turonian, Coniacian and Santonian
age (see Table 2). From the general outlines of the specimens figured on Stoliczka's
PI. 7, figs. 15--17, it is only possible to say that they resemble some variants within
the species Kingena pentangulata (Woodward) from the British Upper Senonian.

Kingena granulifera Stoliczka
1872 Kingena granulifera Stoliczka : 26, pi. 7, figs. 8-12.

Emended description. Elongate-oval to subpentagonal in general outline.
Some specimens show a wider hinge-line than others and tend towards less acute
anterior constriction, thus giving a subparallel or more rectangular outline with steeply
sloping flanks. This form also develops a shallow but noticeable sulcus, beginning
about midway from umbo to anterior margin and broadening slightly anteriorly.

Lectotype. G.S.I. 1589, in the collections of the Geological Survey, India,
Calcutta.

Dimensions of lectotype. Length 18*20 mm., width 14*20 mm., thickness
8*6o mm.

Horizon and locality. Arryalur group, from Olapaudy, Southern India.

KINGENINAE

Remarks. From the five specimens figured by Stoliczka (pi. 7, figs. 8-12) it is
impossible to estimate the full range of variation within the species but they resemble
a similar series of specimens of Kingena mesemhrina (Etheridge) collected from the
Gingin Chalk of Western Australia and now in the British Museum (Natural History)
nos. BB. 45791-BB. 45795. One of these specimens, BB. 35791, figured here PI. 6,
fig. 8, shows the same subparallel flanks and rectangular general outline as K.
granulifera. The growth-marks on the shell surface are as pronounced or step-like
as those on the figured specimens of K. granulifera and a similar extensive, shallow
sulcus is also noticeable in the brachial valve. Elliott (1952) redescribed Etheridge's
species Magas mesemhrinus and assigned it to the genus Kingena, His illustrations
of the species (pi. i, figs. 1-13) can be compared to the drawings of Stoliczka (1872, pi.
7, figs. 8-12).

It may be that the two species are synonymous, but more material from the Indian
locality is needed before any conclusions as to their relationship can be reached.

? Kingena punjabica Sahni

1939 Kingena punjabica Sahni : ii, pi. i, figs. 4, 5.

Holotype. G.S.I. Type no. 16637, the Geological survey of India, Calcutta.

Dimensions of holotype. Length, 16*5 mm., width, 16-5 mm., thickness, ii mm.

Horizon and locality. The stratigraphical details surrounding this species are
somewhat vague and Sahni gives the horizon as ? Neocomian or Upper Jurassic. It
was collected from an exposure two and-a-half miles southeast of Pezu, Bannu
district, N.W. Frontier Province, Pakistan.

Sahni's figured specimen certainly has external morphological features very
similar to those of some Kingena species, especially from the Albian and Lower
Chalk of England, but, as Sahni states ( : 12), the characteristic pustules or granules
on the shell surface are missing.

Similar morphological features are noted in the genus Belothyris which Smirnova
(i960 : 1 17) described from the Lower Cretaceous of the northwest Caucasus.
Examples of this genus have been collected from beds of Lower Cretaceous age at
Sheik Budin, near Pezu in the Bannu district of Pakistan by the Geological Survey of
Pakistan.

Sahni's K. punjabica was collected from the arenaceous beds just above the massive
limestone at Sheik Budin. According to Spath (1939 : 136) the ammonites in these
beds are Crioceratids similar to those from the Trigonia swarzi beds of Tanzania, i.e.
Upper Neocomian. The genus Belothyris, although originally described from beds of
indefinite age, is represented in beds of Upper Neocomian age in France and Switzer-
land by the species B, pseudojurensis.

If, on the other hand, K. punjabica is in fact a true Kingena, then this would be the
earliest record of a species of Kingena, However, without proper examination of
Sahni's specimen and more details of its brachial loop structure, it is impossible to
say for certain as to what genus this species should be assigned. For the present it is
assigned broadly to the genus Kingena.

THE BRACHIOPOD SUBFAMILY

Kingena tumida Muir-Wood
1930 Kingena tumida Muir-Wood : 34, pi. 5, fig. ya-c.

Holotype. No. 14,459 collections of the Geological Survey of India,

Calcutta and was collected from the Gault of the Samana Range, Punjab.

Remarks. Adequately described by Muir-Wood (1930 : 34) as an acutely bi-
convex or globose internal mould with greater convexity of the pedicle valve than the
brachial valve. No fold or sulcus are developed and lateral commissures are straight
with acutely sloping flanks.

Genus BELOTHYRIS Smirnova, i960

Type-species. Belothyris plana Smirnova, i960

Description. The general outline is rounded-pentagonal to elongate-oval, some-
what cinctiform. Acutely biconvex. Rectmarginate to ligate. The slightly pro-
duced umbo is suberect to just incurved with a small circular foramen. The per-
mesothyridid beak-ridges are distinct. A high median septum extends to about
two-thirds the length of the shell. The shell surface is smooth but the growth-marks
are fairly distinct.

Internal characters. Short divergent dental lamellae in pedicle valve support
deeply inserted hinge-teeth. Dental sockets with extensive inner and outer socket-
ridges. Long shallow hinge- trough giving rise to slender hinge-plates supported by
high septum. Descending branches of brachial loop with long inwardly curving
crural processes which diminish rapidly. The long septum develops straight, con-
necting to low transverse band or conical hood from ascending branches.

Remarks. Smirnova (i960 : 115) suggested that in Belothyris there may have
been an earlier stage in its development where the individuals had a connection
between the descending branches and septum, as in Kingena, Although this is a
perfectly acceptable idea, in view of recent comparison between species of Belothyris
and Kingena for the present research, it seems more likely that Belothyris, Kingena
and Zittelina are products of a common ancestor, possibly from the Middle or Upper
Jurassic, which have developed strong individual characters and have remained
separate but closely related genera.

Range. Belothyris is known only from the Lower Cretaceous.

Belothyris pseudojurensis (Leymerie)

(PI. 9, figs. 1-5, 7)

1842 Terebratula pseudojurensis Leymerie : 12, pi. 15, fig. 5, 6.

1847 Terebratula pseudojurensis d'Orbigny : 74, pi. 505, fig. 11-16.
i86i Terebratula pseudojurensis Leymerie ; de Loriol : 121.

1872 Terebratula (W aldheimia) pseudojurensis Leymerie ; Pictet : 93, pi. 203, figs. 11-15.
1896 Zeilleria pseudojurensis (Leymerie) ; de Loriol : 160, pi. 6, fig. 23.

KINGENINAE

Description. Belothyris, elongate-oval to elongate-pentagonal. Ligate anterior
commissure. The umbo is slightly produced and the distinct beak-ridges are per-
mesothyridid to a moderate to small, circular foramen. The general outline is
cinctiform. Some marginal thickening develops in gerontic stages. In some forms
the growth-lines are not particularly distinct, but probably vary with the type of
preservation.

Neotype. As the original specimens figured by Leymerie have been lost, and all
efforts to obtain a specimen from a French source have failed, a specimen has been
chosen from seven specimens collected from the type locality at MaroUes (Aube),
France and now in the Davidson Collection at the British Museum (Natural History).
It is registered as B. 35033 and is figured here, PI. 9, fig. 3a-c. It is here proposed as
neotype for Leymerie's species pseudojurensis referred here to the genus Belothyris.

Remarks. There are many forms within the Lower Cretaceous which might very
well be assigned to the genus Belothyris but there is a need for more research into the
internal structure of these forms. Some of the cinctiform specimens collected from
the Hauterivian and Valanginian of north Germany and figured by Roemer (1840)
may possibly belong to this genus.

Belothyris pseudojurensis appears to be restricted to the French and Swiss Neocomian
beds within the Hauterivian.

Belothyris nettletonensis sp. nov.

(PI. 9, fig. 6a-c)

Diagnosis. Oval to cinctiform. Umbo short, massive. Foramen small, sub-
circular ; beak-ridges sharp. Deltidial plates exposed. Hinge-line moderately
broad ; interarea fairly extensive.

Holotype. From the Lower Cretaceous, Claxby Ironstone, Nettleton, Lincoln-
shire in the British Museum (Natural History) registered no. BB. 45788. The speci-
men was collected by Dr. Peter Rawson.

Description. Medium sized Belothyris, 14 mm. long, 12 mm. wide and 9 mm. in
thickness. Zeilleriform or cinctiform in general outline. The thin test is smooth
and without prominent growth-marks. Marginal thickening is noticeable in the
more mature specimens. The anterior commissure is ligate.

Remarks. The true range of variation cannot be properly ascertained from the
very few specimens available but, in general, it is a short Belothyris not exhibiting the
same degree of anterior sulcation as B. pseudojurensis from France and Switzerland
which it otherwise resembles.

In England B. nettletonensis has been found only in the top beds of the ironstone
facies of the Claxby Series at Nettleton, but probably occurs at a similar horizon at
Speeton in the Speeton Clay.

Material. The holotype, BB. 45788 and two other specimens in the British
Museum (Natural History) no. BB. 45797-98.

THE BRACHIOPOD SUBFAMILY

Genus ZITTELINA RoUier, 1919

Type-species. Terebratula orbis Quenstedt, 1858.

Diagnosis. Oval-pentagonal to subquadrate ; biconvex, smooth. Rectimargin-
ate to incipiently sulcate. Umo short, massive, suberect to slightly incurved ;
foramen small, circular ; beak-ridges indistinct, permesothyridid. Brachial loop
with double attachment to median septum and expanded band to form kingeniform
hood.

Description. Although the type-species is distinctly pentagonal in general
outline, the full morphological range of this genus compares favourably with that of
Kingena from the Cretaceous. Variation within the species, however, does not tend
towards elongation of the valves, as in Kingena, but towards a broader aspect with
increased width of hinge-line. Faint carination of the posterior part of the pedicle
valve appears to be present in early development of some species, particularly Z.
orbis (Quenst.) Z. impressula (Quenst.) and Z. caeliformis Suess.

Internal characters. The brachial loop, as seen in acid developed specimens from
Wurttemburg, shows the characteristic spines on the dorsal surface of the descending
and ascending branches as seen in Kingena and several genera within the Zeilleriacea
from the Middle and Lower Jurassic.

The conical or funnel-shaped kingeniform hood is well developed from the trans-
verse band of the ascending branches but appears to be more quadrate in transverse
outline according to serial sections of the type-species. Fig. 14.

A pedicle collar is a marked feature of the pedicle umbo and the same callus in-
filling of the umbonal cavity is noted which occurs in Kingena, Belothyris and is also
seen in Waconella gen. nov.

From the transverse serial sections of the type species, Z. orbis. Fig. 14, it is pos-
sible to see the marked similarity between the cardinalia and hood attachment in this
genus and the same structures in Kingena, Fig. 2. The differences between the two
genera appear to be confined to the relative angle of the descending branches to
median septum which is more acute in Zittelina, and the length of the median septum
which, in Kingena, always extends to well over two-thirds the length of the shell, but
remains comparatively short in Zittelina, not extending far beyond the point of
attachment of the brachial loop.

Zittelina orbis (Quenstedt)

(PI. 10, fig. la-c. Text-fig. 14)

1858 Terebratula orbis Quenstedt : 639, pi. 29, fig. 28-38.

1870 Megerlea orbis (Quenstedt) ; Zittel : 219, pi. 41, fig. 18.

1871 Terebratula orbis Quenstedt ; Quenstedt : 400, pi. 49, figs. 59-61, 63-74.

1883 Terebratula {W aldheimia) orbis Quenstedt ; Engel : 188-213, 230, pi. 5, fig. 7.

1887 Kingena orbis (Quenstedt) ; Bukowski : 88.

Lectotype. Here selected, the specimen figured by Quenstedt (1871, pi. 49,
fig. 65) from the Upper Jurassic, Middle White Jura, Wurttemburg, Westphalia,

KINGENINAE

Germany. The specimen is in the Quenstedt Collection at the Institut und Museum
fiir Geologie und Palaontologie, der Universitat, Tubingen, Germany.

Description. Zittelina averaging 13 mm. long, ii mm. wide and 7*9 mm. in
thickness. Distinctly oval-pentagonal in general outline. In early growth-stages,

Fig. 14. Series of sixteen transverse serial sections through the umbo of a specimen of
Zittelina orbis (Quenstedt) from the Upper Jurassic of Wurttemburg, Germany. B.

37717- X 3 -

Fig. 15. Series of nine serial sections through the shell of Frenulina sanguiniolenta (Gmelin)
from off the Philippines, Z.B. 3331 showing similar plan of braccial loop to that of
Kingena and Zittelina but indicating the well formed connecting band of septum to
descending branches in the later stages. X 4.

THE BRACHIOPOD SUBFAMILY

the brachial valve becomes slightly inflated posteriorly and maintains this acute
convexity throughout subsequent growth, while the rest of the valve tends to
flatten, especially towards the margins. Growth-marks are prominent but are not
raised much above the shell surface. The median septum is clearly visible in the
brachial valve. Likewise the dental lamellae are distinct as clear lines at the apex
of the pedicle valve. The suggestion of a pedicle umbo between the dental lamellae
in the pedicle umbo was made by Zittel (1870 : 219) but there is no evidence of such
a structure in the serial sections of the type species.

Remarks. Zittelina orhis is probably confined to the European continent, but the
genus is represented elsewhere. Krumbeck (1905) described two new species and
recognized three of QuenstedTs from the Glandarienkalk (Kimmeridgian) of the
Lebanon. These were triangularis and latifrons, which he described as new, and
gutta, cuhica and orhis of Quenstedt, all of which he assigned to the genus Kingena,

Zittel (1870) described a series of beautifully prepared acid developed specimens
from South Germany as Megerlea. Among these was a specimen of Z. orhis (pi. 41,
fig. 8a, b) showing most of the characters described here.

Family LAQUEIDAE Thomson, 1927
Subfamily LAQUEINAE Thomson, 1927

In the “Treatise Hatai (1965 : H. 845) proposed the subfamily name Laqueinae
for Laqtieus, ranging from the Miocene to Recent seas. This name had already been
used in this context by Thomson (1927 : 256) who proposed it as a subfamily name
for Laqueus and Pictothyris. In view of this oversight, the family name Laqueidae
which Hatai also proposed (Hatai, 1965) should therefore be attributed to Thomson
1927 {nom. transl.)

Genus WACOI^ELLA nov.

Type-species. Terehratula wacoensis Roemer, 1852

Diagnosis. Evenly biconvex, oval-pentagonal to elongate-oval. Test smooth.
Punctae coarse, numerous. Umbo prominent, beak-ridges sharp, interarea exten-
sive ; foramen small, permesothyridid. Deltidial plates conjunct. Anterior com-
missure rectimarginate. No cardinal process developed. Brachial loop, laqueini-
form.

Range. Upper Albian to Lower Cenomanian.

Description. External characters. Distinctly terebratulid in general aspect with
a small foramen. The smooth exterior is often without any obvious trace of growth-
marks, although the mode of preservation may tend to make this character more
obvious. Variation within the species is confined to more elongation of the valves to
produce a more elongate-oval outline. The interarea and deltidial plates are usually
well exposed. Slight labiation of the foramen occurs with gerontic development.

KINGENINAE

Internal characters. Nothing is known about the early loop development stages of
Waconella owing to lack of suitable material. Small or young specimens have been
sectioned and the serial sections have revealed that the connecting band from
septum to descending branches in the small or juvenile individuals is considerabl}^
shorter in proportion to the shell than in the more mature forms. This stage is similar
to the ultimate stage in the loop development of Frenulina.

The adult loop in Waconella is virtually the same as in Laqueus, The transverse
band is broad with a wide central aperture. This aperture is formed by the lateral
processes from the transverse band extending towards the descending branches and
connecting with them. At this same point, the lateral bands from the septum also
connect with the descending branches.

Waconella wacoensis (Roemer)

(PI. 10 figs. 6a-c, ya-c ; Text-fig. i6)

1852 Terehratula wacoensis Roemer : 81, pl. 6, fig. 2 a, b, c.

1857 Terehratula wacoensis Roemer ; Conrad : 147, pl. 3, fig. i.

1857 Terehratula leonensis Conrad : 147, pl. 21, fig. 2a-c.

1901 Terehratula (Kingena) wacoensis Roemer ; Hill : 264, pl. 37, fig. 5, 5a.

1928 Kingena wacoensis (Roemer) ; Adkins : 80.

1948 Kingena wacoensis (Roemer) ; Cooper : 365, pl. 143, figs. 32-34.
i960 Kingena wacoensis (Roemer) ; Perkins : 63, pl. 17, figs. 1-8.

Description. External characters. Waconella averaging 20 mm. long, 19 mm.
wide and 12 mm. in thickness. Subcircular to oval in early growth stages, becoming
more elongate-oval in adult stage. Fairly widely spaced, coarse punctae clearly
visible. A moderately long median septum is developed in the brachial valve and
extends from the apex of the brachial umbo to approximately half the length of the
shell. The umbo is slightly produced and is slightly carinate in some individuals.
The hinge-line varies from narrow to moderately wide. The beak-ridges are short,
fairly distinct and permesothyridid. The foramen is small and circular. A fairly
short interarea is reminiscent of that observed in Laqueus.

Internal characters. In transverse serial sections through the shell, the posterior
portion of the umbonal cavity in the pedicle valve is seen to be infilled with callus
which unites the dental lamellae and attaches them to the floor of the valve. Similar
infilling of this cavity has been seen in Kingena, Zittelina and some Recent dallinidae,
such as Macandrevia.

The cardinalia are in many ways similar to those of Kingena, having a broad, fairly
shallow hinge-trough, well marked inner and outer socket-ridges and strong, elongate-
triangular hinge-plates supported by a strong, high median septum. The hinge-
trough flattens anteriorly, giving rise to two ventrally concave plates which lengthen
to form long crural processes and the descending branches of the loop. Proceeding
anteriorly for a short distance, the descending branches are deflected dorsally and
connect with a narrow band from the septum which runs almost horizontal to the
floor of the valve. At this point, the connecting band connects or intersects with
lateral processes produced from the transverse band of the ascending branches.

THE BRACHIOPOD SUBFAMILY

The descending branches, therefore, do not close in to meet the median septum, as in
Kingena, but connect with the septum by means of the interconnecting band, as in
Laqueus, The mature loop develops further down the shell or more anteriorly than
it does in Kingena where the loop develops maturity at the posterior end of the shell .

Fig. i6. Series of twenty-two serial sections through Waconella wacoensis (Roemer),
Duck Creek Formation, Texas, showing similar loop and connecting bands to those of
Laqueus californicus (Koch) as seen in Fig. 17. BB. 5718. X4.

KINGENINAE

Fig, 17. Fifteen serial sections through the shell of Laqueus californiciis (Koch) from
Recent seas, for comparison with sections of Waconella wacoensis (Roemer) in Fig. 16.
Z.B. 3332. X 2 approx.

THE BRACHIOPOD SUBFAMILY

Remarks. Wacondla wacoensis has been recorded from practically every
locality where the Washita strata are exposed. It has been recorded from many
localities in central and western Texas, occurring in the Comanchean Series from the
Edwards and Georgetown Formations, from the Grayson Marl, Del Rio Shale, Main
Street limestone. Pawpaw Shale, Weno Marl, Denton Marl, Fort Worth Limestone,
Duck Creek Formation and the Kiamichi Formation. In terms of European strati-
graphy, it covers beds from Upper Albian to Lower Cenomanian age. (see Table
3 : 40). It abounds in beds of this age in Texas and is frequently found at the same
horizon in Cretaceous deposits in Mexico.

CONCLUSIONS

The present geological range of the Dallinidae is considered to be from the Upper
Trias to Recent seas, but there are many more genera which may belong to this
family still to be thoroughly investigated.

In 1959, Dagis described the genus Aulacothyropsis from the Upper Trias of Siberia
and figured a series of transverse sections of the type species A, rejiexa (Bittner).
These sections show characters which are remarkably similar to those of the Creta-
ceous genus Kingena and also to the Recent genus Frenulina. It may be possible to
trace the lineage of these forms in the course of time and throw some more light on
their evolution. In the meantime, however, there are many intermediate forms
still to be investigated. Some of these are the terebrateUacean genera within the
Middle and Upper Jurassic which are known only from mature adult specimens and
very often from their external morphology alone,.

The attainment of a kingeniform loop may be expected to occur more than once in
the family Kingenidae, for the mechanism of loop-development and evolution
suggested by Elliott (1953) explains a similar phenomenon in the superfamily
Terebratellacea generally, in which the Kingenidae belongs. Thus the future
investigaton of the Jurassic forms mentioned above may clarify the relationship,
if any, of Aulacothyropsis to Kingena.

The genus Rugitela described by Muir-Wood (1936) from the Fuller's Earth Rock
of Somerset and now known to have a more extensive range from Middle Jurassic to
Lower Cretaceous, shows signs of early loop attachment to septum, possibly
dallinoid stages, which are in need of further investigation. Likewise, the genus
Tamar ella described by Owen (1965) from the Upper Aptian of southern England
requires more detailed research on early stages in view of its possible relationship to
Psilothyris, which Cooper described from the Lower Cretaceous of Arizona, before its
proper place in the classification can be established. Similarly, the genus Digonella,
described by Muir-Wood (1934) from the Bathonian, may also have early dallinoid
stages as suggested by that author in her classic work.

All this depends very much on the discovery of very young forms which might
show early loop developmental stages. Such specimens are difficult to find and are
not often recognized even when they have been collected. Perhaps with more
painstaking and more critical observation of small specimens already in collections
it will be possible in time to fill the existing gaps in our knowledge of this fascinating
group of brachiopoda.

KINGENINAE

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E. F. Owen, M. Phil.

Department of Paleontology
British Museum (Natural History)
London, S.W.7

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